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puhusfjed under the patronage of the 
Secretary of State for India. 

EDETBl) Br LT.-COL. K. B. S. SEWELL, C.I.E., Sc.D., E.E.S., l.M.S. iret.). 



B. L. BHATIA, D.Sc, F.Z.S., F.R.M.S., 

1^ FEB 9 1939 t^ 


November 29, 1938. 






Author's Pkbfacb v 

■Glossary of Technical Terms ix 

Systematic Index xv 

Introduction : 

Position of Sporozoa in the Animal Kingdom . . 1 

General Organization and Structure 4 

Phylogeny and Classification 10 

Study of the Group in India 13 

Distribution 17 

(i) List of Parasites and their Hosts .... 18 

(ii) List of Hosts and their Parasites .... 27 

Technique 37 

Systematic Description or Species 50 

Appendix 374 

Addenda 377 

Bibliography 390 

Alphabetical Index 491 



The present volume, dealing with Sporozoa, follows the same 
plan as the author's volume on Ciliophora, published two years 
ago. The group Sporozoa includes numerous organisms, 
all of which are parasitic in hosts belonging to a number 
of different phyla of the Animal Kingdom. Several of these 
organisms are responsible for producing disease in man and 
domestic animals, and consequently come within the ken of 
the medical and veterinary workers : but there are numerous 
others which parasitize animals such as earthworms, poly- 
chsetes, crustaceans, insects, myriopods, arachnids, moUuscs, 
fishes, amphibians, reptiles, birds, etc., and, though causing 
damage to various tissues of the host, are on the whole tolerated 
by them . To the student of Protozoa all are equally interesting 
and it is his province to study them as animals, and name and 
classify them. The medical and veterinary workers will 
find in this volume the correct zoological position and nomen- 
clature of the pathogenic organisms they study. With 
regard to the nomenclature, the synonyms recorded under 
each species will serve to indicate the extent to which opinion 
has in the past varied with regard to the systematic position 
of the particular organisms. In a few cases the reader may 
feel disposed to differ from me, but in such cases he will find 
the argument or the authorities on which I have relied 
clearly stated, and if, after considering them, the reader still 
differs, he is welcome to do so. 

The most approved and up-to-date system of classification 
has been followed, and in the Identification Tables of Families 
I have included those families which are at present not known 

FEB -9 


from India. The 320 species described, in this volume are- 
but a small fraction of the total kno^^^n from other parts of 
the world. All the species that are as yet known from India, 
Bm-ma, and Ceylon have been brought together, but a large 
number still await discovery. The geographical distribution 
of the parasites usually follows that of their hosts. Lists 
of Parasites and their Hosts, and of Hosts and their Parasites 
have been given in the Introduction and the Appendix, 
and it is hoped that these ^vill be of use to those looking- 
for the parasites in particular hosts. It may be pointed out, 
however, that the discovery of a parasite in a host not recorded 
in the list does not mean that the organism is new to science. 
It may aheady have been recorded from the same host or in 
some other host in other parts of the world. Keeping this 
in view, the discoverer should not rely too much on this volume, 
but should carefully explore the monographs and original 
papers to which the Bibliography, given at the end of the volume , 
will furnish a guide. The literature on the subject of Sporozoa 
is very vast, and it has been possible to give a selection onty. 
For the convenience of the reader the Bibliography has been 
divided into a number of sections corresponding to the sub- 
classes or orders, and if a reference is not found under " Text- 
books and Sporozoa in General " it should be looked for in 
the section of the Bibliography relating to the particular 

The species of each genus have been arranged in alphabetical 
order, except in a few cases where the species occurrmg in 
man or some other group of hosts have been brought together. 
In the synonymies, given under each species, references to 
all the records from India, Burma, and Ceylon have been 
included, and a j mark prefixed to all such references. A selec- 
tion of other references which are considered important or 
useful is also given. 

In the Introduction I have given a short account of the 
general organization and structure, and discussed the phylo- 
geny and classification of the group. It seems obvious that 
Sporozoa should preferably be divided into two classes, 
viz., Sporozoa s. str. and Amoebosporidia, but the two classes- 


cannot be arranged in a subphylum, as the former are evolved 
from the Flagellata and the latter from the Sarcodina : so in 
deference to the usage among English authors, the group i& 
described as a single class and is subdivided into a number 
of independent subclasses. A short account of the principal 
methods employed in the study of Sporozoa has also been 

A volume such as this is bound to incorporate very largely 
the work of others, and my grateful acknowledgements are 
due to all those whose works have been drawn upon, and 
especially to Minchin, Wenyon, Hartmann, Reichenow, Calkins, 
Hegner, and Kudo. Where available, figures have been given 
for the species dealt with. A certain number of these are 
taken from my own work, but a large number have been 
borrowed with the kind permission of the authors or publishers 
concerned. Mj^^ thanks are due to the editors and publishers 
of journals and text-books who have given permission to 
reproduce the figures, and due acknowledgement is made in 
every case by giving the name of the author from whom the 
figure has been copied. 

My special thanks are due to Dr. B. Prashad, Director of 
the Zoological Survey of India, for special facilities given to 
me on the occasion of several visits to Calcutta to consult 
the literature in the splendid library maintained by the 
Zoological Survey of India ; and also to Dr. S. L. Hora for 
his help in getting some figures copied under his supervision 
by the artists working under him. Finally, I have to offer 
my most grateful thanks to the Editor, Lieut. -Colonel R. B. 
Seymour SeweU, C.I.E., F.R.S., for a thorough and critical 
revision of the text, and for generous help and guidance during 
the production of this work. 


Hotu Singh Road, Lahore, 
July, 1938. 


Acephaline. — A gregariiie not possessing an epimerite at any stage 

of its life -history. 
Acnidosporidia. — Term sometimes used to include Sarcosporidia and 

Actinoinyxidia. — Cnidosporidia with large spores, trivalved membrane, 

and three distinctly visible polar capsules. 
Adeleidea. — Coccidia in which the gametocytes are dissimilar in size 

and are associated with each other dm-ing the later part of trophic 

Agamete. — An agamic spore or product of asexual reproduction. 
Agamogony. — ^Asexual or agamic reproduction by equal, omequal or 

multiple division. 
Agamont. — An asexual individual reproducing without conjugation or 

Amitotic division. — Direct division of the nucleus unaccompanied by 

the formation of a spindle of threads. 
Amoebosporidia. — Term used by Hartmann to denote a separate class 

of Protozoa, in which are included Cnidosporidia, Sarcosporidia, 

and Haplosporidia. 
Amcebula stage. — ^The sporoplasm which, by amoeboid movements, has 

left the spore membrane, a stage leading up to the schizont. 
Anisogamy. — Conjugation between dissimilar gametes. 
Anterior end. — The end of an organism which is habitually forward 

in locomotion. In Cnidosporidia the end of the spore from which 

the polar filament becomes extruded through the foramen. If 

the two extremities are dissunilar in form, the anterior end is 

usually more or less attenuate. 
Association. — A group formed by the attachment of two or more 

Autogamy.- — Fusion of the two daughter nuclei to form a zygote or 


Biassociative.- — Referring to an association of two sporonts attached 

by imlike ends. 
Binary fusion. — A mode of reproduction in which the division of the 

nucleus into two is followed by the division of the cell. 
Budding. — The process of maequal fission, resulting in the formation 

of daughter organisms, which show a simplified structure when 

first formed. 

FEB " ^ ^-^'-'^ 


Cephaline. — A gregarine possessing an epimerite at some stage in its 

Cephalont. — Young gregarine with an epimerite. 

Chitinous. — Corresponding in appearance and character to chitin, 

the horny material which forms the protective covering of insects 

and other Arthropoda. 

Gnidosporidia. — Sporozoa in which the spore is provided with one or 
more thread-capsules. 

Coccidia. — Telosporidia in which the mature trophozoite is intra- 
cellular and small, the zygote is non-motile, and the sporozoites are 
developed within a spore. 

Coccidiomorpha. — -Term used by Dofiein to include Coccidia and 
Hsemospor idia . 

Ccelozoic. — Parasites that live in the Imnen of the alimentary canal 
or other cavities in the body of the host. 

Conjugation. — Union of two organisms leading to reproduction by germs 
or spores. 

Cortical. — Relating to the external layer of an organism. 

Cyst. — Impervious membrane surroim.duig an organism or a pair of 
associated sporonts at the beginning of reproduction. It is an 
adaptation to a change of hosts in parasitic forms. 

Cytoplasm. — The protoplasm of the cell-body as contrasted with that 
of the nucleus. 

Cytozoic- — Parasites that are lodged inside a cell. 

Deutomerite. — The portion of a cephaline gregarine behind the septum,, 
by which it is separated from the protomerite. 

Dicystid. — A gregarine in which the trophozoite is divided into two 
parts only, that is, an epimerite is present, but the rest of the body 
is not divided into a protomerite and a deutomerite. 

Disporoblastic. — Producing two sporoblasts. 

Disporous. — ^Producmg two spores. 

Ectoparasitic. — Having the nature of an external parasite. 

Ectoplasm.. — The outer zone of the body of unicellular organisms, 

comprising the epicyte, sarcocyte, and myocyte. 
Eimeridea. — Coccidia in which the gametocytes are similar in size and 

develop independently of one another. 
Encystment. — The phenomenon of becoming motionless and excreting 

a membranous cyst. 
Endogenous or internal budding. — Formation of buds in the interior 

of the cytoplasm of the parent. 
Endoparasitlc. — Havmg the nature of an internal parasite. 
Endoplasyn. — The inner or granular zone of the body of unicellular 

organisms, lying within the ectoplasm. 
Endospore. — The inner covering of a spore. 
Epicyte. — The thm, fragile, external layer of the ectoplasm. 
Epimerite. — The temporary, or rarely permanent, structure at the 

anterior end of a gregariiae by which the young gregaruae is attached 

to the host-cell. It is epicytal in origin. 
Epispore. — The outer covering of a spore. 


Eugreqarinaria. — Gregarinida not showing schizogony. 
Exogenous or external buddhig. — -Formation of buds from the external 
surface of the body. 

Fission. — Division of the nucleus followed by a division of the cell-body. 

Gamete. — Specialized cells destined to meet and fuse in conjugation. 

Gametocyst. — The cyst formed round two associated sporonts or gameto- 

Ganietocyte. — The mother -cell which gives rise to a number of gametes. 

Gamogony.- — -The process of prodiiction of gametocytes or gametes by 
a gamont. 

Gamont. — An individual destined to form gametes ; also kiiown as 

Glohidia. — Sarcosporidia in which the spores are fusiform and cysts 
occur exclusively in the intestinal submucosa. 

Golgi apparatus. — A cytoplasmic inclusion which shows a tendency to 
clunip together in masses or to form a network in the neighbour- 
hood of the nucleus. 

Gregarinida. — Telosporidia in which the matiu-e trophozoite is extra- 
cellular and large, the zygote is non-motile, and sporozoites are 
developed within a spore. 

Gyninospore. — Naked germ or protoplasmic body, formed by sporulation, 
which is not enclosed in a protective envelope. 

H'xmosporidia. — Telosporidia in -which the mature trophozoite is 
intracellular and small, the zygote motile, and sporozoites are without 
an envelope. 

Hsemosporidiidea. — Term used by Wenyon to denote Hsemosporidia 
which form pigment in the red blood-cells. 

Haplocyta. — Gregarines in which the trophozoite is not divided by 
an ectoplasmic septum. 

Haplosporidia. — Sporozoa characterized by possession of large spores, 
and a simple type of development. 

Helicosporidia. — Cnidosporidia with small barrel-shaped spores, con- 
taining a thick filament coiled beneath the spore-membrane and 
three sporoplasms. 

Heteropolaridea. — Haplocite, producing sporocysts with dissimilar poles- 

Histozoic. — Parasites that occur in the spaces between groups of cells. 

Holozoic. — Animals which are entirely dependent for food on other 
organisms, which they capture, devour and digest. 

Homopolaridea. — Haplocyta producing sporozysts with similar poles. 

Isogametes. — Gametes which are similar in shape and size. 
Isogamy. — Conjugation between similar gametes. 

Karyogamy. — Union of two gametes whose nuclei undergo intermingling. 
Karyosome. — A chromatic mass surroimded by plastin and contaiiaed 
within the nucleus. 

Longitudinal striations.- — The very delicate ridges on the outside of the 


Macrogamete. — The larger or inactive gamete in anisogamous conjuga- 
Macrogametocyte. — The mother-cell of the macrogamete. 
Meront. — A schizont of a microsporidian. 

Merozoite. — A product of asexual reproduction or schizogony. 
Metabolic. — Changeable in form ; polymorphic. 

Metagamogony. — The process of zygotic or post-conjugation reprodviction. 
Metamorphic. — Changeable in form. 

Microgamete. — The smaller or active gamete in anisogamous conjuga- 

Microgatnetocytes. — The mother-cell of microgametes. 

Microsporidia. — Cnidosporidia with small spores, membrane in one 
piece, and one or, rarely, two polar filaments that are mvisible 
in vivo. 

Microsporoblastic. — Producing a variable number of sporoblasts. 

Microsporous. — Producing a variable number of spores. 

Mitochondria. — Minute cytoplasmic inclusions, of a lipoidal nature, 

occurring in the form of spherical granules or rod-shaped or 

crescentic bodies. 
Mitotic. — Indirect division of the nucleus, which is accompanied by the 

formation of a spindle of threads. 
Monocystid. — Gregarines in which the body of the trophozoite consists 

of a single part, i. e., is not divided by a septum. 
Monosporoblastic. — Developmg into a single sporoblast. 
Monosporous. — -Developing into a single spore. 
Multinucleate. — Possessing many nuclei. 
Multiple fission. — ^A mode of reproduction in which the division of the 

nucleus is not immediately followed by the division of the cell, 

but, after repeated nuclear division, the cell divides into as many 

parts as there are nuclei. 
Myocyte. — The ectoplasmic layer consisting of the myonemes. 
Myoneme. — Specialized muscle-like fibrils which cause the contraction 

of the whole or a part of the body. They are embedded m the 

periphery of the endocyte and form a network around the organism. 
Myxosporidia.- — Cnidosporidia with large spores, bivalved membrane, 

with two or four polar capsules visible tn vivo. 

Neosporidia. — Term used by Schaudinn to include Cnidosporidia, 
Sarcosporidia, and Haplosporidia. The common features are 
that the life of an individtial does not come to an end when repro- 
duction takes place, but that reproduction continues throughout 
the trophic phase, the sporoblasts being carried about by the 
more or less active parent organism, which may ultmiately become 
a large mass of spores. 

Nucleolus. — An exceedmgly minute, more solid particle developed 
singly or in varying number within the nucleus of an anunal or 
vegetable cell. Its homologue among the Protozoa is generally 
referred to as Endoplastule or Karyosome. 

Nucleus. — More densely granular body within the substance of an animal 
or vegetable cell. 

Octosporoblastic. — Producing eight sporoblasts. 

Octosporous. — Producmg eight spores. 

Octozoic spore. — A spore containing eight sporozoites. 


Oocyst. — -A cyst containing the conjugated gametes. 
Ookinete. — The motile zygote in Hsemosporidia. 

Pansporoblast. — -An enclosed area in a myxosporidian trophozoite in 

which two sporoblasts become differentiated. The term is also 

used to designate in general a grown-up sporont of the polysporous 

genera in which two or many sporoblasts are formed. 
Parasite. — An organism living in or upon the body of another organism 

and dependent for its existence on that organism or a limited 

groiip of organisms. 
Piroplasmidea. — Hsemosporidia which do not form pigment in the 

red blood-cells. 
Planont. — -The stage between free amoebula and schizont stages, which 

are found in the alimentary canal or body-cavity of the host soon 

after the spore gerixiinates. 
Plasmodium. — Multinucleated cell formed by the repeated divisions 

of the nucleus. 
Plasmotomy. — Cleavage of a multinucleated body into two or more 

multinucleate parts. 
Polar capsule. — A sac in which the polar filament is coiled, a structure 

characteristic of a cnidosporidian spore. 
Polar filainent. — ^A fine and long filament coiled in the polar capsule, 

which is extruded under suitable stimulation. 
Polycystid. — -Referring to gregarines possessing a septum which divides 

the trophozoite into regions ; also known as " septate." 
Polymorphic. — Exhibiting a diversity of form. 
Polysporoblastic. — Producing numerous sporoblasts. 
Polysporous. — Producing many spores. 
Posterior end. — -The end of an organism which is habitually behind 

in locomotion. In Cnidosporidia the end of the spore opposite 

to the anterior ; it is usually more or less rounded if the two ends are 


Primite. — The first individual in an association of two or more sporonts. 
Protomerite. — -The portion of a septate gregarine which precedes the 

Protoplasm. — The physical basis of life, or elementary formative matter 
of all living organisms. 

Protozoa. — Animals in which the body is not divided into cells. 

Pseudocyst. — ^The residual protoplasm which, after the spores are 
separated, acquires a membranous wall, swells until the true cyst- 
wall bursts, and allows the extrusion of the ripe spores. 

Sarcocyte. — The middle layer of the ectoplasm. 

Sarcosporidia. — Sporozoa in which cysts form long rod-like masses, 

and the spores are crescentic. 
Satellite. — -Any sporont in an association which is attached behind the 

primite. Generally there is one, but sometimes several are attached 

linearly, one behind the other, or in a cluster to the posterior 

end of the primite. 

Schizogony. — ^Asexual or agamic reproduction by equal, unequal or 
multiple division. 

Schizogregarinaria. — -Gregarinida showing schizogony. 
Schizont. — That stage which is about to divide into a number of parts 
called merozoites. 


Schizontocyte. — A special form of agamont which breaks up into a 

number of agamete-forming centres by multiple division. 
Septata. — Gregarines in which the trophozoite is divided by an ecto- 

plasmic septum. 
Septum. — The thin layer of sarcocyte which separates the protomerite 

from the deutomerite. 
Siliceous. — -Partaking of the nature and qualities of silica ; composed 

of flint. 
Spore. — The body into which the zygote develops after the acquisition 

of a resistant outer coating. 
Spore-duct. — A tubular growth from the cyst, through which the spores 

are extruded when ripe. 
Sporoblast. — A product of the initial reproduction of the zygote, 

including both capsule and contents ; a cell which develops 

directly into a spore. 
Sporocyst. — The covering or coverings of the spore. Sometimes used 

in the same sense as a spore. 
JSporogony. — The development of spores from the sporont. 
Sporont. — An adult sporozoan which is destined to form gametes or 

to give rise to sporoblasts. 
Sporoplasm. — The protoplasmic mass inside the spore ; the sporozoite 

of a Cnidosporidian spore. 
Sporozoa. — A class of the Protozoa in which spore-formation is the 

prevailing mode of reproduction. 
Sporozoite. — Each of the small falciform bodies which are released when 

the spore-wall is absorbed. 
Sporulation or multiple fission. — -Mode of reproduction in which the 

repeated division of the nucleus is followed by the splitting of the 

organism into as many parts as the nuclei. 
Syngamy. — Sexual union or conjugation involving a complete fusion 

of two gametes. 
Synkaryon. — The combination nucleus resulting from the fusion of 

two nuclei derived from two individuals. 
Syzygy. — Linear chains formed by two or more organisms attached 

end to end. 

Telosporidia. — -Sporozoa with trophic and reproductive phases typically 
distinct. Term used by Schaudinn to include Gregarinida, 
Coccidia, and Hsemosporidia. 

Tetrasporoblastic. — Producing four sporoblasts. 

Tetrasporous. — Producing four spores. 

Tetrazoic spore. — A spore containing four sporozoites. 

Trophozoite. — -The young feeding and growing parasite. 

Vacuolate. — Having a number of clear spaces or vacuoles. 
Vermicular. — -Resembling a worm in shape. 

Zygote. — The cell resulting from the complete fusion of two gametes. 



Class SPOROZOA Leuck 50 


Schaud 50 


Schneid. em . Doflein . 52 

I. Suborder Eugeegarin- 

ABiA Doflein 53 

1. Legion Haplocyta -Lan^^. . 54 
1. Tribe Homopolarldea 

Bhatia 54 

1. Fam. MoNOCYSTiB^ St. em. 

Bhatia 55 

1. Gen. Monocystis St. em. 55 

Hesse <Sb Cognetti .... 55 

1. beddardi Ghosh 56 

2. bengalensis Ghosh .... 57 

3. Uoidi Ghosh 58 

4. pheretimi Bh. d: Gh.. . . 58 

2. Gen. Apolocystis Cogrnefi* . . 59 

1. matthaii {Bh. cfc Set.). . 59 

3. Gen. Nematocystis i?esse . . 60 

1. hessei Bh. ds Ch 60 

2. lumbricoides Hesse ... 62 

3. plurikaryosomata Bh. 

<hCh 62 

4. stephensoni Bh. cfc Set. 63 

5. vermicularis Hesse .... 64 


Bhatia 65 

1. Gen.Rhynchocystisfl'esse. 65 

1. awatii Bh. <h Set 66 

2. cognettii Bh. cb Ch. . . 67 

3. mamillata Bh. cfc Set. . 67 
S. Fam. Stomatophorid^ 

Bhatia 68 

1. Gen. StomatophoraDrzew. 

em. Hesse, (t Bhatia . 68 

1. bulbifera Bh. <b Set. . . 69 

2. coronata (Hesse) 69 

3. diadema Hesse 71 

4. primitiva Bh. ds Set. . . 377 

1. cambolopsisse Chak. . . 93 

2. rawiiGhak. da Mit 95 

Irtb-y 1939 

4. Fam. Zygocystid^ Bhatia. 72 

1. Gen. Extremocystis Setna 72 

1. dendrostomi Setna .... 72 

5. Fam. Aikinetocystid^ 

Bhatia 73 

1. Gen. Aikinetocystis (?aies. 73 
1. singularis Gates 73 

2. Gen. Nellocystis Gates ... 75 
1. birmanica Gates 75 

6. Fam. Diplocystid^ Bhatia 76 

1. Gen. Lankesteria Mingaz. 11 

1. culicis (Ross) 77 

2. mackiei (Shortt ds 79 

Swami.) 79 

3. tripteroidesi, sp. nov. . . 81 

2. Tribe Heteropolaridea 

Bhatia 82 


Lecudinid^ Kamni 
em. Reichen 

1. Gen. Lecudina Mingaz. 

1. brasili Ganap. db Aiy 

2. eunicse Bh. c& Set. . 

3. lysidicae Bh. c& Set. 

2. Gen. Bhatiella Setna 
1. morphysse Setna . 

3. Gen. Ferraria Setna . 
1. cornucephali Setna 






Incert^ sedis. 

1. Gen. Dirhynchocystis Cogf- 

netti 87 

1. globosa (Bh. d: Ch.) . . 87 

2. Gen. Grayallia Setna .... 88 
1. quadrispina Setna .... 88 

2. Legion Septata Lank. ... 89 

1. Fam. Stenophorid^ Leg. 

d: Dub 91 

1. Gen. Stenophora Labbe . . 91 

1. ellipsoidi Chakravarti . 91 

2. khagendrse Ray ...... 92 

2. Gen. Hyalosporina Chak. . 92 




2. Fam. Grbgaeinid^e Labhe. 96 

1. Gen. Leidyana Watson. . . 97 

1. gryllorum (Cuenot) ... 97 

2. xylocopje Bh. &: Set. . . 98 

2. Gen. Gregarina Dufour . 98 

1. aciculata, sp. nov 99 

2. cornwalli, sp. nov 100 

3. oviceps Diesing 101 

3. Gen. Caulocephalus Bh. <h 

Set 102 

1. crenata Bh. S Set. ... 102 

4. Gen. Protomagalhsensia 

Pinto 103 

1. attenuata Set. ds Bh. . . 104 

5. Gen. Hirmocystis Leger. . 106 
1. parapeneopsisi^'ef. (f?-B/i. 106 

6. Ulivina Mingaz 380 

1. eunicEe Bh. tfc Set 380 

7. Deuteromera Bh. (t Set.. . 381 
1. cleava Bh. <h Set 381 

8. Contortiocorpa Bh. do Set. 383 
1. prashadi Bh. cfc Set 383 

3. Fam. Actinocephalid^ 

Leger 107 

1. Gen. Steinina -Legf. <£■ -Dm6. 107 
1. metaplaxi Pearse 107 

4. Fam. Dactylophorid^ 

Leger 108 

1. Gen. Grebneckiella, nom. 

nov 108 

1. navillse [Mit. ct Ghak.). 108 

5. Fam. Monoductid^ Pay 62 

Ghak 109 

1. Gen. Monodtictvis Pay <£• 

Ghak 109 

1. lunatus -Ra?/ cfc Ghak. 109 

II. Suborder Schizogkegar- 

INARIA Leger Ill 

Fam. Selenidiid^ Brasil . . . 384 

1. Gen. Selenidium Giard. . . 384 

1. amphinomi i?/i. cfc iSe^.. . 385 

II. Order COCCIDIA Leitcter;;. 113 

I. Suborder Adeleidea 

Leger 113 

1. Fam. Adeleid^ Leger .... 115 

1. Gen. Adelea A. Schneid. 115 
1. pachelabrse de Mello . . 116 

2. Gen. Adelina Hesse 116 

1. schellacki Pay <£• Das- 

Qup 116 


2. Fam. H^mogregarinid^ 

Leger 117 

1. Gen. Hasmogregarina 

Danil 119 

1. berestneffi Cas<. da Will. 119 

2. caxA\\Q\ Samh . ds Selig . . 121 

3. hankini Simond 121 

4. laverani Simond 122 

5. leschenaultii Robertson. 123 

6. magna (Grassi d; 124 


7. malabarica de Mello . . 124 

8. raesnili Simond 127 

9. mirabilis Gast. S Will.. 127 

10. najse Laveran 128 

11. nicorise Gast. d> Will... 129 

12. nucleobisecans Shortt. . 131 

13. phythonis [Billet) 132 

14. rara Lav. ofc Mesn 133 

15. rodriguesi de Mello <£• 

others 134 

16. stepanowiana Lav. db 

Mesn 134 

17. testudinis Lav. ds Natt.- 

Lar 135 

18. thomsoni Minchin .... 136 

19. thyrsoidese de Mello <& 

Vales 138 

20. triedri Pobertson 138 

21. vittatfe Robertson 139 

22. xavieri de Mello 140 

23. sp. Plimmer 142 

24. sp. Simond 142 

25. sp. Dobell 142 

26. sp. Pobertson 142 

27. sp. Plim,mer 143 

28. sp. Simond 143 

29. sp. Dobell 143 

30. sp. Dobell 143 

31. sp. Patton 143 

32. sp. Patton 143 

33. sp. Patton 144 

34. sp. Plimmer 144 

35. sp. Simond 144 

36. sp. Simond 144 

37. sp. Robertson 144 

38. sp. {Billet) 145 

39. sp. Patton 145 

2. Gen. Hepatozoon Miller . 145 

1. adiei Hoare 146 

2. canis {James) 147 

3. felis domestici {Patton). 149 

4. funambuli {Patton) .... 149 

5. gerbilli {Ghristophers) . 150 

6. leporis {Patton) 153 

7. muris {Balfour) 153 

8. sp. Donovan 154 


3. Gen. Karyolysus Labbe . . 154 
1. jorgei de Mello cfc de 

Meyr 155 

II. Suborder Eimeridea 

Leger 156 

1. Fam. EiMERiiD^ Leger ... 158 

1 . Subfam. Ceyptosporidin^ 

Poche 159 

1. Gen. Lankesterella Labbe 

1. minima (Chaussat) .... 159 

2. monilis (Labbe) 162 

2. Subfam. Cyclosporins 

Wenyon 162 

1. Gen. Isosjiova A.Schneider 162 

1. belli Wenyon 163 

2. bigemina (Stiles) 165 

3. calotesi, sp. nov 167 

4. felis Wenyon 168 

5. knowlesi Ray <&; Das- 

Gup 170 

6. minuta Mit. cfc Das- 

Gup. 170 

7. rivolta (Grassi) 171 

8. wenyoni Ray cfc Das- 

Gup 172 

9. sp. Cooper <t Gulati ... 173 

3. Subfam. Eimeriins Wen- 

yon 173 

1. Gen. Eimeria J.. /S'c7me?rfe?' 173 

1. arloingi (Marotel) 386 

2. clupearima (T/?.e7o/ian). . 177 

3. colimibiE 3Iit. dk Das- 

Gup 178 

4. cylindrica Wilson 387 

5. faurei (Mous.d; Marat.) 179 

6. flaviviridis Set. <h Bana. 179 

7. gupti, nom. nov 386 

[for cylindrica Ray & 

Das-Gup 179] 

8. Yiav^odom. Set. (b Bana . 181 

9. hemidactyli Knoivles db 

Das-Gup 182 

10. kermorganti (Simond). 183 

11. knowlesi Bhatia 184 

12. koormae Das-Gupta . . . 184 

13. labbeana Pinto 387 

14. laminata Ray 185 

15. legeri (Si^nond) 186 

16. mitraria (Lav. & Mesn.) 187 

17. najse Ray tf? Das-Gup. . 188 
18 piscatori Ray <£■ Das- 
Gup 189 

19. sardinse (Thelohan) 189 

20. smithi Yak. <£■ Gal 191 

21. sonthweWi Halawani . . . 193 


22. vvassilewskyi Rasteg 

23. yakimovi Rasteg. . 

24. ziirni (Rivolta) ... 

25. sp. Cooper db Gulati 

26. sp. Setna <b Bana 
Setna c& Bana 
Setna db Bana 
Setna d; Bana 
Setna (h Bana 

sp. Setna <&i Bana 
sp. Setna & Bana 
sp. Setna & Bana 
sp. Setna dh Bana 

sp. Ross 

Gen. Wenyonella Hoare 
1. hoarei Ray 



4. Subfam. Barrotjssins 

(Wenyon) 199 

1. Gen. Pythonella Ray <&; 

Das-Gup 199 

1. bengalensis Ray <&; Das- 
Gup 199 

5. Subfam. Aggregating 

Reichenow (em. Hoare) . . 199 

1. Gen. Aggregata Frenzel . . 200 

1. sp. Setna db Bhatia .... 202 

Incerts sedis. 
1. Gen. Toxoplasma Nic. db 

Mane 202 

1. butasturis de Mello . . . 203 

2. canis U. Mello 204 

3. columbse Yak. <&; Kohl- 

Yak 204 

4. eimiculi Splendore 205 

5. fulicse de Mello 206 

6. sp. Adie 207 

7. sp. Plimmer 207 

8. pyrogenes Castellani . . 207 


Danil 209 

1. Fam. H.a;MOPROTEiDG 

Doflein 210 

1. Gen. Hsemoproteus itrwse. 211 

1. antigonis de Mello 212 

2. asturis de Mello 212 

3. cerehneisi, sp. nov 212 

4. columbaa Celli S San 

Pel 213 

5. coracias de Mello ct 

Afonso 217 

6. corvi (de Melio) 218 

7. danilewskyi (Grassi <&; 

Feletti) 219 





8. glaucidii de Mello 221 

9. gymnorbidis de Mello . 221 

10. herodiadis f/e ilf eZZo 221 

11. kopki (de Mello) 222 

12. machlolophi de Mello . . 223 

13. metchnikovi (Simond) . 224 

14. orioli de Mello 225 

1.5. raymundi de Mello da 

JRaim 225 

16. rileyi Malkani 226 

17. simondi (CasZ. c& T^4ZZ.). 227 

18. sturni de Mello .' . 228 

19. upupse de Mello 228 

20. wenyoni de Mello c& 

others 229 

21. sp. Scott 230 

22. sp. Scott 230 

23. sp. Donovan 230 

24. sp. Scott 230 

25. sp. Cast, cfc Will 230 

26. sp. Plimmer 230 

27. sp. Plimmer 231 

28. sp. Scott 231 

29. sp. Donovan 231 

30. sp. Scott 231 

21. sp. Plim')ner 231 

32. sp. de Mello 231 

33. sp. Knowles 232 

34. segithinse de Mello 232 

35. anthi de Mello 232 

36. bram«3 de Mello 232 

37. centropi de Mello 233 

38. dicruri de Mello 233 

39. elani de Mello 234 

40. gallinulse de Mello 234 

41. halcyonis de Mello 234 

42. halcyonis var. fuscse de 

Mello 234 

43. lanii de Mello 235 

44. otocompsse de Mello . . 236 

45. pastoris de Mello 236 

46. platalese de Mello 237 

47. tephrodornis rfe MeZZo . . 237 

48. iheveicery CIS de Mello . . 237 

49. thereiceyrcis var. zey- 

lonica de Mello 238 

2. Gen. Leucocytozoon 

Danil 238 

1. ardeolse de Mello 239 

2. chloropsidis de Mello . . 239 

3. coracise de Mello <fc 

Afonso 240 

4. enriquesi de Mello .... 241 

5. melloi, sp. nov 241 

6. molpastis de Mello .... 242 

7. sp. Donovan 242 

8. sp. Donovan 242 

9. sp. Scott 242 

1 0. sp. Knowles 243 


11. sp. Plimmer 243 

12. sp. Plim?ner 243 

13. sp. knowles 243 

14. sp. de Mello 243 

15. sp. de Mello 244 

2. Fam. Plasmodiid^ Mesnil. 244 

1. Gen. Proteosoma Labbe. . 247 

1. centropi (de Mello) .... 249 

2. chloropsidis (de Mello). 250 

3. columbse (Garini) 250 

4. gallinxilse (de Mello . . . 251 

5. herodiadis (de Mello) . . 252 

6. heroni (Basu) 252 

7. moruony (de Mello cfc 

de Sd) 253 

8. passeritae de Mello ds 

da Pons 388 

9. pr9sco-s:(Grassi(i; Feletti) 253 

10. sp. Scott 257 

11. sp. Donovan 257 

12. sp. Plimmer 258 

13. sp. Plimmer 258 

14. sp. Plimmer 258 

15. sp. Plimmer 258 

16. sp. Scott 258 

17. sp. Scott 258 

18. sp. Plimmer 259 

19. sp. Plimmer 259 

20. sp. Plimmer 259 

21. sp. Plimmer 259 

2. Gen. Laverania Grassi <fc 

Feletti 260 

1. raalavis?, Grassi d; Feletti 260 

3. Gen. Plasmodium iJfarc/ii. 

cfc Celli 270 

1. vivax (Grassi d; Feletti). 271 

2. malarise (Laveran) 273 

3. cynomolgi Mayer 281 

4. inui Halb. ds Prow. . . . 283 

5. linowlesi Sinton dh Mul- 

ligan 284 

6. kochi Laveran 285 

7. ptheci Halb. da Prow. . 286 

8. semnopitheci Knowles . 287 

9. bubalis Sheather 289 

10. canis Cast, d; Chalm. . . 290 

11. equi Cast, d; Chalm. . . 290 

1 2 . mackiei de Mello SdeSd 291 

13. narayani de Mello d) 

Dias 291 

14. pteropi Breinl 292 

15. ratufe Donovon 293 

16. tyrio de Mello d' others. 293 

3. Fam. Theileriid^ cZw ToiZ . 294 
1. Gen. Theileria Beit., 

Franca, c& Borg 295 

1. parva (Thciler) '. 295 




2. mutans (Theiler) 298 

3. sp. Sen 6a Srinivasan. . 299 

4. CQ\\\i(Cast.<&; Ghalm.).. 300 

0. hirci Dschun. <Ss Urod. . . 300 

4. ,Fam. Babesiid^ Poche . . . 301 
1. Gen. 'Babesia, Starcovici . . 302 

1. bigemina (Smith c& 

Kilb.) 303 

2. bovis (Babes) 306 

3. caballi (Nuttall cfc 

Strickl.) 307 

4. canis (Piana ds Galliv.). 307 

5. equi (Laveran) 312 

6. felis Davis 313 

7. gibsoni (Patton) 314 

8. motasi Wenyon 316 

9. iiinense (Yakinioff) ... 317 

10. sergenti Wenyon 317 

11. soricis (Christophers) . . . 318 

12. taylori (Saruar) 301 

13. tropicus (Ling, cfc Jenn.) 318 

14. sp. Patton .'. . 320 

1.5. sp. Plimmer 320 

16. sp. Patton 320 

Incebt^ sedis. 
Pam. Anaplasmid^ Nietz, 

Alex., ds du Toit 320 

1. Gen. Grahamella BrwTOpi. 320 

1. canis (Kikuth) 322 

2. muris Carini 323 

3. sp. de Mello 323 

2. Gen. Pai'aplasixia Seidelin 324 
1. Mello da others . . 324 

3. Gen. Anaplasma Theiler 325 

1. sp. de Mello cfc others. . . 325 

2. sp. Knowles 326 

3. marginale Theiler 388 

. 4. Gen. Bertarellia Carini. . 326 

1. calotis de Mello db de 

Meyr 326 

2. carinii de Mey relies ... 389 

3. sp. de Mello tfc de Meyr. 327 


Doflein 328 


Biitschli 330 

I. Suborder Eurysporea 

Kudo 333 

Fam. Cbratomyxid^ Doflein. 334 
1. Gen. Cer atomy xa Thelo- 

han 334 

1 . gobiodesi Chakravarty . . 334 

2. hilsse Chakravarty .... 335 

II. Suborder Sph^rosporea 

Kudo 335 

1. Fam. Chloromyxid^ The- 

lohan 335 

] . Gen. Chloromyxum 

Mingazzini 335 

1. amphipnovi Ray 336 

2. Fam. Sph^rosporid^ 

Davis 336 

1. Gen. Sphserospora Thelo- 

han 336 

1. sp. Southw. d; Prashad 336 

III. Suborder Platysporea 

Kudo 336 

1. Fam. Myxidiid^ T/ie'Zo/ian . 337 

1. Gen. Myxidium Biitschli. 337 

1. danilewskyi Laveran. . . 337 

2. glossogobii Chakravarty. 337 

3. mackiei Bosanquet .... 338 

4. sp. Ray 339 

2. Gen. Zschokkella Auer- 

bach 339 

1. prashadi Ray 339 

2. Fam. Myxobolid^ Thelo- 

han 340 

1. Gen. My xobolus Bitoc/iZ/ . 340 

1. calhasui Chakravarty .. . 340 

2. norigalse Chakravarty . . 341 

3. nodularis Southw. <fc 

Prashad 341 

2. Gen. Thelohanellus Kudo 342 

1. rohitse (Southw. tfc Pra- 

shad) 342 

2. seni (Southw. d; Pra- 

shad) 344 

3. Gen. Henneguya Thelohan 344 

1. ophiocephali Chakra- 

varty 345 

2. otolithus Ganapati .... 345 


Stole 346 

1. Fam. Haploactinomyxid^ 

Granata 346 

2. Fam. Euactinomyxid.e 

Granata 346 


Balbiani 346 

1. Suborder Monocnidea 

Leg. <&; Hesse 348 

Fam. NosEMATiD^ Labbe .... 349 
1. Gen. Nosema Ndgeli em. 

Perez 349 

1. adiei (Christophers) . . . 349 




2. bombycis Ndgeli 351 

3. ctenoeephali Kudo .... 354 
2. Gen. Thelohania ffe^megruy 356 

1. indica Kudo 356 

2. legeri Hesse 357 

3. obscura Kudo 359 

Gen. et sp. incert. Ross .... 359 

II. Suborder Dicnidea ie'^/. 

cfc Hesse 360 

Fam. Telomyxid^ Leg. ds 

Hesse 360 


Kudo 360 


DIA Butschli 361 


s. str., Babudieri 362 

Fam. SAECOCYSTiD>gE Babu- 
dieri 363 

1. Gen. Sarcocystis Z^an- 

kester 363 

1. blanchar di Do/ein .... 363^ 

2. lindemanni (Rivolta) . . 365 

II. Order GLOBIDIA Babu- 
dieri 367 

Fam. Globidid^ Babudieri . . . 367 

1. Gen. Globidium Flesch .. 367 

1. fusiformis Hassan .... 367 

2; sp. Gilruth <fc Bull. . . 368 


DIA Liihe 369 

Doubtful Protozoa. 

Gen. Rhinosporidium Min. 

cfc Fanth 370 

1. seeberi ( H^ermc&e) 370 

2. sp. Vasudevan 372 

3. equi Zschokke 372 

4. sp. Rao 373 


Position of Sporozoa in the Animal Kingdom. 

Sporozoa are a class of parasitic organisms belonging to the 
phylum Protozoa. In the Protozoa the body usually 
consists of a single undivided mass of protoplasm, and these 
organisms are consequently described as unicellular or non- 
cellular. In other classes of the Protozoa the animals, 
which may be free-hving or parasitic, move by characteristic 
organs of locomotion such as pseudopodia, flagella or ciha : 
but the class Sporozoa includes organisms which are entirely 
parasitic, and as a rule are incapable of free locomotion. 
Some of them, when immature, move about by means of 
pseudopodia, but they never possess ciha or fiagelJa. All 
are characterized by producing spores. 

Our knowledge of the Sporozoa commenced just over 
a hundred years ago. In 1826 Dufour gave the first detailed 
account of an organism, which he afterwards, in 1828, named 
Gregarina. He found it in the ahmentary canal of several 
species of Coleoptera, and later in the gut of the earwig. 
Both he and Siebold, who investigated the genus m 1837 and 
1839, failed to recognize it as a Protozoan. Schleiden, Henle, 
and others recognized the unicellular nature of Gregarina, 
but regarded it as an overgrown plant-cell, and it was only 
as late as 1848 that Kolliker demonstrated the Protozoan 
nature of the genus, described the formation of pseudonavicellse, 
and inferred that they represented one method of propagation, 
a view that was confirmed the same year by Stein. The 
complete fife- cycle of a Gregarine was worked out much 
later by Siedlecki (1899). 

In 1839 Hake pubhshed the earhest account of a Coccidian. 
He described and figured the oocysts of Eimeria stiedse of the 
rabbit's liver, but did not mterpret them as of parasitic origin. 
Important observations on a Coccidian were pubhshed by 
Kloss (1855), who described the organism now known as 
Klossia helicina in the renal organ of the snail. Eimer (1870) 
described parasites of the gut and the fiver of various animals, 
including man, which have since been referred to the genera 


P^B - 9 1939 


Eimeria and Isospora. Pfeiffer (1892) was the first to describe 
the theory of alternate generations, and Schaudinn and 
Siedlecki (1897) showed that there are two kinds of gametes 
in Goccidium, and that the macro- and microgametes conjugate 
to form zygotes. Later (1898) Siedlecki pubHshed an account 
of the sexual cycle of the Coccidian Aggregata, and in 1899 
gave a detailed account of the entire ]ife-history of Adelea. 
This series of classical researches led up to the pubHcation of 
the description of the complete hfe- cycle of Goccidium schubergi 
by Schaudinn (1900). 

The H^MOSPOEiDiA, or blood-inhabiting Sporozoa, were 
discovered as late as the eighties of the last century. Laveran 
observed the malarial parasites in the human blood in 1880, 
and pubhshed a description in the following year. He observed 
the amoeboid, rosette, sphere, crescent and the flagellate stages, 
but did not determine the relationship of the various stages, 
nor did he recognize the animal nature of the parasites. The 
genus Plasmodium was instituted by Marchiafava and Celh 
(1895). Further discoveries leading to the elucidation of the 
complete hfe-cycle of the malarial parasites were made by 
Metchnikoff (1887), Golgi (1889), Danilewsky (1891), Manson 
(1894, 1896), MacCallum (1897, 1898), Ross (1898, 1899), 
Grassi (1898), and others, and will be referred to elsewhere in 
this volume. Smith and Kilborne (1893) demonstrated that 
Babesia, the cause of Texas fever of cattle, was transmitted 
from host to host by ticks, and were thus the first to show the 
important part played by insects in the transmission of para- 
sitic Protozoa. 

We owe the term Sporozoa to Leuckart (1879), who intro- 
duced it to include the Gregarinida and the Coccidia. The 
H^MOSPORiDiA were included later, when the study of the 
complete hfe-cycle showed the close resemblance of various 
stages with those of the Coccidia. Besides these three well 
recognized groups, there are other groups of parasites, found 
in Fishes, Arthropods, etc., which, on account of their forming 
spores, came to be included in the Sporozoa. In a large 
majority of these, known as the Cnidosporidia, the spore 
contains a coiled filament inside a polar capsule. Schaudinn 
(1900) divided the Sporozoa into two subclasses — (i) the 
Telosporidia, to include the Gregarinida, Coccidia, and 
H^MOSPORiDiA ; and (ii) the Neosporidia, to include the 
Cnidosporidia, Sarcosporidia, and Haplosporidia. The 
members of both these subclasses produce resistant spores, 
but according to many authors the affinities between the two 
are not sufficiently close to justify their inclusion in the same 
class. Hartmann (1907) estabHshed two classes, for which 
he employed Schaudinn's names — Telosporidia and Neo- 
sporidia. Later, Prowazek and JoUos (1922) and Hartmann 


(1923-5) designated the two classes as Sporozoa and 
Amcebosporidia, and something can be said in favour of 
this system of classification. Wenyon (1926) also divided 
the entire group into two classes, and named them as 
Sporozoa and Cnidosporidia, leaving the Sarcosporidia and 
Haplosporidia as parasites of undetermined position. 
Theoretically speaking, there is no difficulty in restricting 
the term Sporozoa to the original connotation which Leuckart 
gave it, but in practice it would seem impossible so to restrict 
it, unless a new term like Sporogbnea is coined for a sub- 
phylum to include the two classes. In view of the term 
Sporozoa having become fixed in its extended usage, and the 
phylogeny of the two groups being altogether hypothetical, 
Reichenow (1929, 1935) divided the class Sporozoa into foxu" 
distmct subclasses, viz., Telosporidia, Cnidosporidia, 
Sarcosporidia, and Haplosporidia. Similarly Kudo (1931) 
and Calkins (1933) divide the class Sporozoa into three 
subclasses, Telosporidia, Cnidosporidia, and Acnido- 
SPORIDIA, combining in the last the Sarcosporidia and Haplo- 
sporidia. The balance of opinion at the moment does not 
seem to favour the recognition of only two classes, but, if 
this were done, they would be named Sporozoa {sensu stricto) 
and Amcebosporidia. For practical convenience, and in 
deference to general practice, I shaU follow Reichenow (1929, 
1935) in dividing the class Sporozoa (sensu lato) into four 

The phylum PROTOZOA may thus be divided as follows :— 
A. Subphylum Plasmodroma Doflein, 1901, emended. 

Movement effected by pseudopodia or fiagella, and syngamy 
takes place, in all known cases, by the complete fusion of gametes. 

I. Class Mastigophoba Diesing, 1865. 

The predominating phase flagellate, locomotion being effected 
by filamentous whip-like structures called flagella. The body 
may be corticate or non-corticate. 

II. Class Rhizopoda von Siebold, 1845 ( = Sae,codina Hertwig & 
Lesser, 1874). 
The predominating phase amoeboid, locomotion being effected 
by temporary extensions of the body called pseudopodia. 
The body is non-corticate, i. e., has no tough limiting mem- 
brane or cuticle. 

III. Class Sporozoa Leuckart, 1879. 

Exclusively parasitic forms which lack definite organs of loco- 
motion. Reproduction takes place by spore-formation. 

(a) 1. Subclass Telosporidia Schaudinn, 1900 ( = Sporozoa 

sensu stricto). 

Trophozoite becomes full grown before reproduction 
begins ; spore simple, with one to several sporozoites 
or without resistant envelope ; the sporozoite a gre- 
garinula ; asexual and sexual reproduction alternate 



(b) Neospobidia Schaudinn, 1900 (=AMfEBOSPOBiDiA Hart- 
Trophozoites may begin to form spores when still growing 
or even quite young ; the sporoblasts are formed by a 
process of internal gemination ; the sporozoite is an 
amoebula ; and the hfe-cycle is passed in as ingle host. 
This group includes the remaining three subclasses. 

2. Subclass Cnidosporidia Doflein, 1901. 

Spore with polar filament which is tj^pically coiled within 
a polar capsule. 

3. Subclass Sakcospoeidia Balbiani, 1882. 

Spores crescentic, without thread capsules. Cysts forming 
long rod-like masses. Parasites of striped muscles of 

4. Subclass Haplospoeidia Caullery & Mesnil, 1905. 

Spores large, contauiing a single volimainous nucleus ; no 
thread- capsules. Type of development simple. 

B. Subphylum Ciliophoea Doflein, 1901. 
Movement effected by cilia. 
IV. Class CiLiATA Perty, 1852. 

Organisms bear cilia throughout Hfe. 

V. Class SucTOEiA Claparede & Lachmann ( = Tentaculifera Huxley, 
A cinetaria Lankester ) . 
Ciliated in the young stages, but later usually attach themselves 
to other objects, lose their cilia, and develop knobbed tentacles 
which serve as sucking tubes. 

General Organization and Structure. 

This volume deals with the class Sporozoa as defined above, 
and I give below a brief survey of the general organization 
and structure of the organisms included in this group, so as 
to give the reader a general idea of the group and to introduce 
him to the principal technical terms employed in the 
description of the forms. 

Modes of Life. — All Sporozoa are obhgatory parasites, and 
there are no free-hving forms among them. Parasitic forms 
are also found among other classes of the Protozoa, but 
these may be regarded as free-living forms which have been 
introduced casually into the body of their host and have 
become adapted to a parasitic mode of life. In Sporozoa 
the dependance has reached an extreme Hmit, and the parasites 
have no existence apart from the hosts in which they are found 
to occur. The transference of the parasite from one host to 
another is effected by means of spores, which may be defined 
as resistant seed-hke bodies, containing one or more sporozoites 
or germs and protected by a firm envelope or capsule. The 
spore is a contrivance to enable the parasite to withstand 
the vicissitudes of the outside world until they pass again 
into the body of a suitable host. Once inside the body of 
the new host, the spore germinates and a fresh infection is 


started. The mode of transfer of the spores is usually con- 
tamuiative, that is to say, the spores are ingested with the food 
that has been contaminated with faeces in which the spores 
have been passed. In some forms, however, where the parasite 
passes part of its hfe- cycle in the blood of one host and part 
in the body of a widely different host, the sporozoite is not 
enclosed within a definite membrane or cyst, and the method 
of transfer is inoculative. 

The method of nutrition is by osmosis only. The organism 
does not possess pseudopodia, flagella or ciHa for the purpose 
of food-capture. Where the organism is amoeboid, its pseudo- 
podia are for the purpose of increasing the extent of the body- 
surface for absorption, rather than for the ingestion of solid 
particles of food. The parasite may invade the cells (cytozoic), 
or spaces between the cells {histozoic),ov may hve in the lumen 
of the ahmentary canal or other cavities in the body of the host 
(coelozoic). The food material absorbed from the host will 
thus be dissolved cytoplasm, tissue fluid, body fluid, or digested 
food material from the ahmentary canal of the host. The 
Spoeozoa are parasitic in animals of almost every phylum 
from Protozoa to Chordata, and while many of them have 
come to be tolerated, others are responsible for causing 
deadly disease and heavy mortahty among the hosts. In 
accordance with thek widely varied habitat, they show 
manifold adaptations suiting them to the highly speciahzed 
conditions of their existence. 

Form and Structure. — The Sporozoa show a more or less 
comphcated hfe-history, consisting of various stages. The 
starting point is the minute germ or sporozoite, which may have 
one of two forms. In Amcebosporidia it is a minute amoeboid 
organism termed an amoebula ; in the Telosporidia it is 
more definite in form, being a rod-like or sickle-shaped body 
(" falciform body ") which is capable of twisting or bending 
movements and progresses by ghding, and is described as 
a gregarmula. The sporozoite, after being set free in the body 
of the new host, nourishes itself and grows, often to a relatively 
large size, at the expense of the host. This is the trophic 
phase, and the organism during this phase is described as the 
trophozoite. The trophozoite absorbs nourishment in the 
fluid state, and does not exhibit any organs of locomotion, 
ingestion or digestion ; neither food- vacuoles nor contractile 
vacuoles are present. The parasite has usually a fixed form 
with definite contours, being limited externally by a cuticle 
of greater or less thickness, and may either grow inside a host 
cell or be attached to a cell of the host by a special organ of 
fixation known as an epimerite. The ectoplasm is in certain 
groups differentiated into three layers, known as epicyte, 
sarcocyte, and myocyte, the latter being composed of myonemes 


or contractile filaments which bring about contortions of the 
body. The sarcocyte may run inward as one or more septa 
and thus divide the body into a number of apparent segments : 
in the cephahne Gregarines (fig. 30) the body is thus seen to 
consist of three segments, known respectively as the epimerite, 
protomerite, and deutomerite. Sometimes the organisms may 
attach themselves to each other and form clusters or linear 
chains, termed syzygies. The endoplasm is usually granular, 
and contains sorted-up food material in various forms. The 
nucleus is usually very large, spherical, and vesicular in type, 
with one or more distinct karyosomes. 

Reproduction. — Like other parasites, Spokozoa possess the 
power of prohfic multiphcation as a necessary adaptation for 
the maintenance of the species. Both asexual and sexual 
methods are known in all groups. Asexual reproduction may 
be by binary fission (as in Babesia), by multiple fission (as in 
CocciDiA and H^mospokidia), or by budding, which may be 
exogenous (as in Myxosporidia) or endogenous (as in certain 
Schizogregarines). The trophozoite when it is about to enter 
on asexual reproduction is known as a schizont (or agamont), 
the process being termed schizogony (or agamogony) and 
the resulting products of division being termed merozoites. 
Sexual reproduction [syngamy) is by isogamous or anisogamous 
conjugation or sometimes by the fusion of sister individuals 
derived by fission of the same parent cell or nucleus {autogamy). 
Trophozoites which associate with one another for the purpose 
of sexual reproduction are designated as sporonts or gametocytes, 
which may differ from one another and are then known as 
microgametocytes and macrogametocytes, according as they give 
rise to males or microgametes and females or macrogametes. 

The fusion of two gametes results in the formation of 
a zygote which, when activelj^ motile and vermiform (as 
in H^mosporidia), is generally knoAvn as an ookinete. 
The zygote maj^ secrete a distinct membrane round itself 
and become a passive spherical body known as an oocyst. 
Cellular division of the protoplasm within the oocyst gives rise 
to the formation of a number of sporoblasts, each of which 
becomes surrounded by a wall, the sporocysf, and is then 
known as a spore. Inside a spore are developed a smaller 
or larger number of minute germs or sporozoites which are 
the infective bodies, and start the cycle again. These various 
stages, starting from the zygote and leading to the formation 
of the spores and sporozoites, constitute sporogony. Asexual 
and sexual methods of reproduction regularly alternate in the 
life-cycle of the majority of forms, and compHcated hfe-histories 
result. In such cases schizogony takes place usually in the body 
of one host and gametogony and sporogony in the body of 
an animal belonging to a widely different group. The former 


is described as an intermediate host, and the latter, in which 
the sexual process is perfected, as the final or definitive host. 

Life-history. — The hfe-histories of different groups vary 
a good deal, but there is always some resemblance to a common 
fundamental type. The hfe-history of the Coccidium Eimeria 
stiedse (Lindem.) (fig. 1), occurring in the liver and intestine 

Fig. 1.— Life-cycle of a tjrpical coccidium, Eimeria stiedse (Lindem.). 
1-5, sporogony; 6-15, schizogony; 17-20, gametogenesis ; 
21-23, fertilization. (After Reich). 

of the rabbit, may be described as typical. The sporozoite (6) 
escapes from the spore in the intestine of the host, and, gKding 
like a minute Gregarine, bores its way into the epithehal 
cell (7) and becomes a trophozoite (8). The latter feeds at 
the expense of the cytoplasm of the host -cell and grows into 


a large spherical schizont with a vesicular nucleus containing 
a large karyosome. The nucleus of the schizont divides 
repeatedly until sixteen or thirty-two daughter nuclei are 
produced (9, 10), and the schizont breaks up into an equal 
number of merozoites (11). The merozoite may attack another 
epithelial cell and undergo schizogony again, or, on entering 
a fresh epithehal ceU, may produce a set of only four mero- 
zoites (13-15), which differ from the others in possessing 
a flagellum (16). The flagellated merozoites penetrate 
other epithelial cells and become either macrogametocytes 
(17 a) or microgametocytes (17 b). The macrogametocyte 
grows into a large oval body containing many chromatinoid 
and plastinoid granules, which gather round the periphery 
to form a membrane (17 a-20 a), and thus gives rise to a 
single macrogamete (21). The microgametocyte becomes 
large and spherical, and within it are formed a large number 
of biflagellated microgametes (17 6-20 6). FertiHzation (21) 
takes place either in an epithehal cell or in the lumen of the 
intestine. The resulting zygote (22, 23) passes out of the 
intestine as an oocyst in the faeces (1). The protoplasm 
within the oocyst contracts into a spherical mass (2), and 
then divides into four spherical sporoblasts (3), which 
become ovoidal (4), secrete a sporocyst, and become spores, 
part of the cytoplasm not being used up in the process. Within 
each spore two sporozoites develop (5), again leaving a small 
amount of residual protoplasm. Fresh infection takes place 
through the food becoming contaminated with faeces containing 
these spores. 

While the above life-history may be taken as typical, 
considerable variations are met with in different groups. 
In the majority of Sporozoa the hosts are of the same species, 
and infection is brought about by eating contaminated food ; 
but in many forms there is an alternation of hosts, schizogony 
taking place in an animal of one group and gametogenesis and 
sporogony in a host belonging to an entirely different group. 
Thus in the case of Aggregata asexual reproduction takes place 
in a Crustacean (crab) and sexual reproduction in a Cephalopod 
(cuttle-fish). Among the H^mosporidia schizogony takes 
place in the blood of a Vertebrate (man, other mammals, or 
various birds) , while sexual reproduction, followed by sporogony, 
occurs in an Insect (mosquito). In such cases spore -capsules 
are not formed, as the sporozoites are introduced cUrectly 
into the blood by the mosquito. 

In the Eugregarmes the asexual cycle is entirely wantmg, 
the sporozoite developing directly into a gametocyte. Two 
individuals lie side by side, and an envelope or gametocyst is 
secreted enclosing both individuals. Each individual then forms 


a large number of gametes, those from one individual fusing 
with those from the other, and a corresponding number of 
zygotes are formed. Within the membrane the zygotes 
develop into spores, each dividing to form a definite number 
of sporozoites. 

Among the Amcebosporidia (=Nbosporidia) the life-cycle 
is less compUcated, and sexual dimorphism and change of hosts 
are not met with. Reproduction takes place more or less 
continuously throughout the trophic phase, the organism 
ultimately becoming a huge mass of spores. Spore-formation is 
also on an entirely different plan from that in the Telosporidia, 
and does not usually result from the divisions of a zygote. 
The life-histories among the subclasses Cnidosporidia, 
Sarcosporidia, and Haplosporidia differ considerably, and 
it is impossible to attempt a generahzed account : reference 
must therefore be made to the detailed accounts given under 
these different subclasses. Among the Cnidosporidia both 
schizogony and sporogony are met with. Schizogony is 
carried out by binary or multiple fission, by budding, or by the 
cleavage of the multinucleated plasmodium into two or more 
multmucleate parts (plasmotomy) . Isogamous, anisogamous, 
and autogamous reproduction have been reported in a number 
of forms, and the zygote becomes the sporont, in which 
one to many spores become differentiated. The spores are of 
a unique structure. Each spore possesses one to four polar 
capsules, each containmg a coiled polar filament, and one to 
many sporoplasms. The mode of development of these spores 
differs in different cases. In the orders Myxosporidia and 
AcTiNOMYXiDiA Several cells appear during the process. 
These cells give rise to one or more sporoplasms or generative 
cells, capsulogenous cells, and the spore-membrane. In 
the order Microsporidia the amoeboid sporopiasm undergoes 
schizogonic multiphcation, and the schizonts later become 
sporonts, each producing the characteristic number of spores. 

The Sarcosporidia produce long rod-like masses of spores 
among the muscle-fibres, which they parasitize. The spores 
are crescentic in outhne and do not contain polar capsules. 
Only portions of the full hfe-cycle are known. 

The Haplosporidia show a comparatively simple Hfe-cycle. 
The spores are spherical or elhpsoidal, with a single large 
nucleus and no polar capsule. The amoebula grows, and by the 
repeated division of the nucleus forms a plasmodium. The 
Plasmodium may divide (plasmotomy) or may produce mero- 
zoites (schizogony) or form spores. The spores arise either 
from sporoblasts, each of which gives rise to a single spore, 
or from pansporoblasts, which give rise to a number of 


Phylogeny and Classification. 

All parasitic forms are assumed to have been evolved from 
free-living ancestors. The problem natm-aUy presents itself 
as to what must have been the nature of the ancestral forms of 
the Sporozoa and what is their relationship to the remaining 
classes of the Protozoa. The Ciliophora are at once 
excluded, as they are far too speciahzed, and the Sporozoa 
do not show any special relationship with them. Two rival 
theories of Sporozoan ancestry have been put forward in the 
past, one claiming their descent from the Rhizopoda, the other 
from the Mastigophora. The Rhizopoda and the Mastigo- 
PHORA are themselves hnked together. Many Rhizopoda 
show flagellated stages in then- life -cycle, and there are many 
Mastigophora which are amoeboid. Most authorities agree 
with Awerinzew (1910) that an amoeboflagellate type represents 
the primitive stock of the Protozoa, which gave rise to all 
existing groups, becoming differentiated into Rhizopoda on 
the one hand and Mastigophora on the other. Biitschh 
(1882) was the first to advance the theory of the Euglenoid 
ancestry of Sporozoa, according to which a typical Flagellate 
would become adapted first to a saprophytic and then to 
a parasitic mode of life, and thus lose the special organs of 
locomotion, nutrition, etc. As remarked by Minchin (1903), 
" an Euglena or Astasia deprived in this way of flagellum, 
mouth, chromatophores, stigma, and vacuoles, nutritive 
or contractile, would be practically indistinguishable from 
a simple Gregarine." The euglenoid movements of the Gre- 
garines, and of the motile stages of other Sporozoa, such as 
the sporozoites and merozoites of Coccidia, the free stages 
of the HiEMOGREGARiNiDA, and the ookinetes of the IL^mo- 
SPORiDiA, lend support to this view, and additional support 
has been furnished by the discovery of flagellated stages in the 
hfe- cycle in many Sporozoan forms. 

The life-cycles of the Amcebosporidia (=Nbosporidia), 
however, do not lend support to the above-mentioned view. 
They have no Euglenoid phases, do not possess flagella at any 
stage of their life, and are amoeboid throughout their trophic 
phase. Thus the Amcebosporidia give support to the rival 
theory of Rhizopod ancestry. It was doubtless this funda- 
mental difi'erence that led Schaudinn (1900) to divide the 
Sporozoa into two subclasses, viz., Telosporidia and 
Neosporidia ; and led Hartmann (1907) to regard the 
Sporozoa {sensu stricto) and the Amcebosporidia as distinct 
classes. Minchin (1912) expressed the opinion that from such 
forms as Cercomonas arose on the one hand the Rhizopoda 
and their derivatives (Neosporidia) by loss of flagella and 
speciaHzation of the amoeboid form in the adult, and on the 


other the Mastigophora and their derivatives (Telosporidia, 
Infusoria) by speciaHzation of the flagellar apparatus, 
combined with the acquisition of a cortex and loss of amoeboid 
movement. Hartmann (1923-5) and Fantham (1936) have 
also discussed the philogeny of the Sporozoa. 

Taking the Telosporidia (^Sporozoa sensu stricto) first, 
we are struck by the obvious points of difference between the 
Gregarinida and the Coccidiomorpha. In the Grbgarinida 
two sporonts become enveloped in a common cj^st before they 
give rise to gametes, and the union of gametes takes place 
within the cyst. The cyst contains many zygotes, and each 
zygote gives rise to a single spore. In the Coccidiomorpha, 
on the other hand, the gametocytes are more or less widely 
separated from one another when producing the gametes ; the 
female gametocyte remains undivided to form a single macro - 
gamete, which is very much larger than the microgametes ; 
and the zygote undergoes a process of division, giving rise to 
a large number of sporoblasts and sporozoites. The common 
ancestral type may be assumed to have been one in which 
the trophozoites that grew into gametocytes were separated 
from one another and produced their gametes separately, as 
in CocciDiA ; but each gametocyte produced a number of 
gametes which were more or less ahke, as in the Gregarinida. 

As the Gregarinida came to acquire an intercellular trophic 
phase, the sporonts became free and motile, and it was thus 
possible for the gametocytes to associate and encyst together, 
producing their gametes in close proximity. There would 
be neither any difficulty for the male gametes to find the 
female, nor any need for increased speciaHzation of the gametes. 
They would lose even the slight degree of speciaHzation in- 
herited from the ancestral form, with the result that they 
would be similar and would be produced in equal numbers by 
the two gametocytes. 

The CocciDiA, on the other hand, retained an intracellular 
habitat for their trophozoites, and the gametocytes were 
widely separated when producing the gametes. The gametes 
thus had to seek each other, and this led to greater speciaHza- 
tion. The male gametes became very small and very motile, 
and were produced in large numbers ; the female gametocyte 
no longer divided into a number of gametes, but became a single 
macrogamete. After fertilization the suppressed divisions 
of the female gametocyte would take place in the zygote, 
leading to the production of a number of sporoblasts, spores, 
and sporozoites. The spore may be regarded as comparable 
to the encysted zygote of the Flagellata. In the suborder 
Adelbidea, however, the gametocytes acquired the habit of 
association prior to gamete formation, but this led merely 
to a reduction in the number of male gametes produced. 


The H^MOSPORiDiA resemble the Coccidia very closely. 
Their Ufe- cycle can be described in identical terms and the 
points of diflference attributed to their becoming adapted to 
parasitism upon a special kind of cell, viz., the blood- corpuscles : 
the two orders are thus so closely alhed that Doflein (1901) 
was led to consider the two as suborders of a single order, the 
CocciDiOMOEPHA. The H^MOSPORiDiA exhibit an alternation 
of generations, the asexual process or schizogony alternating 
with the sexual process leading to sporogony ; but an essential 
difference from the Coccidia is that there is an alteration 
of hosts, schizogony taking place in the blood or internal 
organs of a Vertebrate, and sporogony in the digestive tract 
or other organs of an Invertebrate. Fertihzation takes place 
in the stomach of the Invertebrate host, which has sucked up 
the gametocytes from the blood of the Vertebrate. The 
zygote, instead of being a motionless body, is a motile vermicule 
(ookinete), which penetrates the wall of the stomach and forms 
its oocyst, which increases in size with the growth of the zygote, 
and only persists while the zygote is producing sporozoites. 
The absence of spores with resistant cysts is due entirely to 
the fact that the parasite is always sheltered within the body 
of one or the other of its two hosts. 

So far as the Amcebosporidia (=:Nbospoeidia) are concerned, 
their affinities are entirely with the Rhizopoda. The body- 
form of the sporozoite and the adult is that of an amoeba, 
and no flagellated stages are known to occur. As remarked 
by Minchin, " the union of the Telospokidia and Neospokidia 
in one class — the Sporozoa — ^is a quite artificial arrangement." 
Practical convenience and common practice alone justify 
their inclusion in one class. 

The Amcebosporidia comprise the three subclasses Cnido- 
sporidia, Sarcosporidia, and Haplosporidia. The Cnido- 
SPORIDIA are a well-defined group and are characterized by the 
spores possessing the polar capsules. Laveran and Mesnil 
(1899) described in the spores of Sarcocystis tenella a striated 
structure representing a polar capsule, and Minchin (1912) 
was led to include Sarcosporidia among the Cnidosporidia. 
Later authorities have shown that among the Sarcosporidia 
there is nothing corresponding to the polar capsule, so the 
Cnidosporidia, Sarcosporidia, and Haplosporidia are best 
regarded as distinct subclasses, having no affinities with one 
another or with the Tblosporidia. 

The Cnidosporidia comprise the orders Myxosporidia, 
AcTiNOMYXiDiA, and MiCROSPORiDiA, to which a fourth 
order has been added by Kudo (1931), under the name of 
Helicosporidia, to include a single species described by 
KeiHn. Dunkerly (1925), discussing the development and 
relationship of the Myxosporidia, pointed out that these, 
like the Volvocace^ among the Flagellates, represent an 


unsuccessful line of advance from the typical Protozoan, 
not reaching, however, the Metazoan type of structure. 
He discussed the origin and relationship of the spore -forming 
nuclei and cells in the pansporoblast, and suggested that 
physiologically the spore of a Myxosporidian is a multicellular 
unit analogous to the Infusoriform embryo of the Mesozoan 
Dicyema, although the Myxospokidia exhibit Rhizopodan 
relationships, while the Mbsozoa are probably derived from 
ciHated ancestors. Although the Myxospokidia do not 
represent a direct link between Protozoa and Mbtazoa, 
they seem to indicate a physiological reason for the origin of 
a soma, as a protective accessory to germ-cells. There is 
a well-marked alternation of generations among the Ccelen- 
TEEATA, as there is among the Myxospokidia, and the occur- 
rence of nematocysts in the former and the polar-capsules in 
the latter is not without significance. 

The Spokozoa are divided in this work into the following 
subclasses, orders, and suborders : — 

I. Subclass Telosportdia Schaudinn. 

I. Order Gregaeinida A. Schneider em. Doflein. 

1. Suborder Etjgregarinaria Doflein. 

1. Legion Haplocyta Lankester. 

2. Legion Septata Lankester. 

2. Suborder Schizogregabinaria Leger. 
II. Order Coccidia Leuckart. 

1. Suborder Adeleidea Leger. 

2. Suborder Eimeridea Leger. 

III. Order HLsimosporidia Danilewsky era. Doflein. 

1. Suborder H^mosporidhdea Wenyon, 

2. Suborder Piroplasmidea Wenyon. 

II. Subclass Cnidosporidia Doflein. 
I. Order Myxosporidia Biitschli. 
II. Order Actinomyxidia Stole. 

III. Order Microsporidia Balbiani. 

IV. Order Helicosporidia Kudo. 

III. Subclass Sarcosporidia Balbiani. 
I. Order Sarcosporidia Babudieri. 

II. Order Globidia Babudieri. 

IV. Subclass Haplosporidia CauUery & Mesnil. 

The further classification into famihes and the genera and 
species dealt with will be seen from the Systematic Index. 

Study of the Group in India. 

Vandyke Carter (1888), Evans (1888), Hehir (1893), and 
Crombie (1894) were the earhest to observe the malarial 
parasites in India. It was, however, Ronald Ross (1895, 1897) 
who observed certain stages of development of the malarial 


parasites in the stomach of the mosquitoes fed on the blood 
of malarial patients, and later (1898) elucidated the life-cycle 
of the malarial parasites of birds and their transmission by 
Culex mosquitoes. In the course of his investigations between 
1895 and 1899 he also noted a number of other parasites of 
mosquitoes. Ross had not received any special protozoological 
or entomological training, and had to coin a new terminology 
for the various phases of the parasites that he observed. 
Since that time India has been the field of the labours of many 
workers on different groups of Sporozoa, and it will be con- 
venient briefly to review their work by deahng with the various 
orders one by one. 

Gregabinida. — Ross (1895) was the first to describe 
a Gregarine parasite from a mosquito, which he referred to as 
Gregarina culicis. Guenther (1914) described a parasite from 
another mosquito in Ceylon, and Mackie (1915) and Swaminath 
(1923) recorded certain organisms from sandflies in Assam 
and Bengal, which were later fuUy described by Short and 
Swaminath (1927). All these organisms are now knoAvn to 
belong to the genus Lankesteria. Cornwall (1915), while study- 
ing the anatomy and hfe-history of Lepisma saccharina (?), 
described some of its Gregarine parasites, which have been 
considered as new species of Gregarina in the present work. 
Ghosh (1923) described some monocystids from the earth- 
worms of Calcutta. 

About the same time I took up the study of the monocystids 
of earthworms in the Punjab, and Bhatia (1924), Bhatia and 
Chatter] ee (1925), and Bhatia and Setna (1926) described 
quite a number of new species, belonging to the genera Mono- 
cystis, Apolocystis, Nematocystis, Stomatophora, Bhynchocystis, 
and Dirhynchocystis, from the earthworms from the Punjab 
and Bombay. Setna (1927) described a remarkable new 
organism under the name Grayallia quadrispiiia from Pheretima 
heterochseta from Bombay. Bhatia and Setna (1924) described 
several cephahne Gregarines from certain Insects, including 
Caulocephalus crenata from a beetle and Leidyana xylocopse 
from the carpenter-bee, the latter being the first Gregarine to 
be recorded from a h3mienopteran host. Later Setna (1931) 
described three new Gregarines, Bhatiella morphysse, Ferraria 
cornucephali, and Extremocystis dendrostomi from certain 
Polychsetes, etc., taken at Port Blair in the Andaman Islands. 
Setna and Bhatia (1934) also described some new Gregarines 
from a prawn from Bombay. 

Gates (1926, 1933) gave a description of two very curious 
parasites from the coelom of certain Burmese earthworms, 
Aikinetocystis singularis and Nellocystis birmanica, which have 
had to be placed in a family by themselves. 

Working at Calcutta, Ray initiated a series of studies on 


Spobozoa from Indian Millipedes, and Ray (1933), Ray and 
Chakravarti (1933), and Chakravarti (1933, 1935, 1936) have 
described Stenophora khagendree, Stenophora ellipsoidi, Mono- 
ductus lunatus, Hyalosporina cambolopsisse, and Hyalosporina 
rayi. Three of these forms are placed in new families by their 

CocciDiA. — Simond in 1901 was the first to pubKsh a series 
of papers on Hsemogregarines from Indian tortoises and the 
gavial, and Laveran and Mesnil (1902) and Laveran and 
Nattan-Larrier (1912) observed Hsemogregarines in certain 
other tortoises. The pioneer work on the HiEMATOZOA in 
Ceylon Avas carried out by Castellani and Willey (1904, 1905), 
Robertson (1908, 1910), and Dobell (1910). James (1905), 
Bentley (1905), Christophers (1905, 1906), and Patton (1908, 
1909) described species of Hsemogregarina and Hepatozoon 
from various mammals, and the sexual cycle of Hepatozoon 
canis was fully worked out by Christophers (1907, 1912). 

Froilano de Mello and his colleagues, working in Portuguese 
India, have pubhshed a number of papers (1915-1937) dealing 
with the Haimogregarines of hzards and Toxoplasmids of 
various birds. De Mello also described (1921) Adelea pache- 
lebrse in a mollusc. 

In the family Eimeriidse many species are now known 
from hosts belonging to all classes of Vertebrates. Cooper 
and Gulati (1926), Cooper (1926, 1927), Sen (1932), and Ware 
(1936) recorded cases of bovine Coccidiosis. Eaiowles and 
Das-Gupta (1931, 1934) studied the Coccidia of the mon- 
goose, cat, and lizard, and Das-Gupta (1934) recorded a case 
of coccidial infection in man. Ray (1935 a, h) described 
Wenyonella hoarei from the squirrel and Eimeria laminata 
from the common toad ; and Ray and M. Das-Gupta (1935) 
described Isospora wenyoni from the toad, and also (1937 a, b) 
described Coccidia from a Hzard and the cobra. Setna and 
Bana (1935 a, b), working in Bombay, described Eimeria 
harpodoni from a fish and Eimeria fiaviviridis from a lizard. 
They have also recorded the occurrence of Coccidia in a 
number of species of marine fish. 

H^MOSPORiDiA. — Ab mentioned already, Vandyke Carter 
(1888), Evans (1888), Hehir (1893), and Crombie (1894) were 
the earhest to observe the malarial parasites in India. But it 
was the memorable work of Ronald Ross (1895, 1897, 1898) 
that laid the foundation of the correct understanding of the 
life-cycle of the malarial parasites in man and birds. He 
succeeded in estabhshing the fact that further development 
of the human malarial parasites takes place in the body of the 
dapple-winged mosquitoes that have fed on the blood of malarial 
patients ; and he elucidated the complete life -cycle of the malarial 


parasites of birds and the part played in their transmission by 
Culex mosquitoes. His discoveries were confirmed by Daniel Is 
(1899, 1900), Stephens and Christophers (1903), and Christo- 
phers (1904). Cornwall (1901), Donovan (1909), and Cragg and 
Naidu (1918) recorded olaservations on the morphology of the 
malarial parasites. In more recent years important contri- 
butions have been made to our knowledge of the morphology 
of these parasites by Acton, Curjel, and Dewey (1921), Knowles 
(1923), Lai (1925), Knowles and Senior White (1927), Clark 
(1927), Hehn (1927), Row (1928, 1930), Knowles, Acton, 
and Das-Gupta (1929), and Knowles (1931). Sinton (1929) 
has published a complete bibliography of the hterature 
deahng with malaria in India. Papers dealing with trans- 
mission by various species of Anophehnes and the seasonal 
incidence of malaria include (among others) those of Stephens 
and Christophers (1902), James (1903), Bentley (1911), James 
and Liston (1912), J. R. & H. Adie (1913), Gill (192.5), James, 
Nicol, and Shute (1927), Carter and Jacocks (1929), Carter 
(1930), Knowles, Senior White, and Das-Gupta (1930), King 
(1931), and Iyengar (1931-4). The distribution of the various 
species of malarial parasites has been dealt with by Christo- 
phers and Sinton (1926), and the distribution of different 
species of Anopheline carriers in important monographs by 
Covell (1927, 1931). 

The malarial parasites of monkeys have been studied by 
Knowles (1919), Donovan (1920), Knowles and Das- Gupta 
(1932, 1934), Smton and Mulhgan (1933), Smton (1934), and 
Mulhgan (1935). Malarial parasites have also been described 
in various other mammals, viz., by Mackie (1914) in a bat, 
Sheather (1919) in the buffalo, de Mello and Paes (1923) 
in the horse, de Mello and colleagues (1928) in the ant-eater, 
and de MeUo (1936) in the otter, and in various birds by de 
MeUo (1935). 

Parasites belonging to the family Hsemoproteidse have been 
described by Castellani and Willey (1904, 1905), Acton and 
Knowles (1914), Alcock (1914), Adie (1915), de Mello and 
colleagues (1917), de MeUo and Raimundo (1934), de Mello 
(1935), and de Mello and Afonso (1935). 

The Piroplasmids have also been studied by a number of 
workers. Lingard and Jennings (1904) were the earhest to 
observe these parasites in various animals. Webb (1906) 
observed them in fox-hounds and Axe (1906) in horses. 
Christophers (1907) described the developmental stages of 
Piroplasma canis in the tick, and Patton (1910) described 
P. gibsoni from the dog and the jackal. Baldrey (1911), 
Gaiger (1911), and Symons and Patton (1912) also studied 
the Piroplasmids. Sinton (1921) described Nuttallia ninense 
from the hedgehog, Rau (1926) and Symons (1926) studied the 


Piroplasmids from hounds, Cooper (1926) from cattle, 
Krishna Iyer (1933) from goats, and Achar and Shrikantiah 
from sheep. Sarwar (1935) described Piroplasma taylori 
from a goat, but I (1936) have shown that this form 
appears to be identical with Theileria hirci. Shortt (1936) 
has re-studied the Hfe-history of Babesia canis in the dog- 

Cnidosporidia. — Very Httle work has been done on the 
Cnidosporidia. The Myxosporidia have been studied by 
Bosanquet (1910), Southwell (1915), Southwell and Prashad 
(1918), and Ray (1933). As regards the Microsporidia, 
Korke (1916) described a species of Nosema parasitic in the 
dog-flea. Mrs. H. A. Adie (1922) observed certain bodies in 
Cimex rotundatus in a Kala-azar infected area in Assam, and 
beheved them to be Leishman-Donovan bodies, but Christo- 
phers (1922) and Short and Swaminath (1922) regarded the 
organism as a species of Nosema, and described it as N. adiei. 
Iyengar (1929) and Kudo (1929) have studied the micro- 
sporidian parasites of Anopheline larvae. 

Sarcospobidia. — Shipley (1904) and Willey, Chalmers, and 
Phillip (1904) described sarcosporidian infection in buffaloes 
in Ceylon ; Chatter jee (1907) described what was probably 
the same species from the heart-muscle of a cow in Calcutta. 
Vasudevan (1927) described a case of sarcosporidian infection 
of man in Madras. Hassan (1935) has described a new species 
of Globidium from a cow. 

Haplosporidia. — Minchin and Fantham (1905) described 
an organism, which they named Rhinosporidium kinealyi, 
from a nasal polyp, and regarded it as belonging to the Haplo- 
SPORiDiA, but Ash worth (1923) has shown that the organism 
is a fungus. Vasudevan (1932) described a case of rhino- 
sporial infection of the fore-arm of a man in Ceylon ; but 
no true Haplosporidia have so far been described from 


The geographical distribution of parasites usually follows 
that of their hosts. The names of the hosts, and the localities 
in which any parasite has been found, are noted after the 
description of the species, and, generally speaking, the same 
parasite may be expected to occur in the same species of hosts 
in other locahties also. The following Usts of (i) parasites 
and their hosts, and (ii) the hosts and their parasites, will, 
it is hoped, be found useful, and indicate at a glance which 
of our commoner animals still remain to be examined for their 

SPOR. c 



(i) List of Parasites and their Hosts *. 

Parasite. Host. Seat. 
Fam. MoNOCYSTiD^. 

Monocystis beddardi .... Eutyphoeus nicholsoni . Seminal vesicles. 

EutyphoRus sp Seminal vesicles. 

Monocystis bengalensis . . Pheretima posthuma . Seminal vesicles. 

Monocystis lloidi Pheretima posthuma . Seminal vesicles. 

Monocystis pheretimi .... Pheretima posthuma . Seminal vesicles. 

Apolocystis matthaii Megascolex trilobatus . Seminal vesicles. 

Nematocystis hessei Pheretima heterochseta . Seminal vesicles, 

Nematocystis lumbricoides Pheretima heterochseta . Seminal vesicles. 

Nematocystis plurikaryo- Allolobophora {Eisenia) Seminal vesicles. 

somxita. foetida. 

Nematocystis stephensoni . Eutyphoeus incommodus Seminal vesicles. 

Nematocystis vermicularis . Pheretima barbadensis . Seminal vesicles. 

Fam. Rhynchocystid^. 

Rhynchocystis awatii .... Pheretima elongata . . . Seminal vesicles. 

Rhynchocystis cognetti . . . Allolobophora caliginosa Seminal vesicles. 

Rhynchocystis mamillata . Pheretima elongata . . . Seminal vesicles. 

Fam. Stomatophorid^. 

Stomatophora bulbifera . . Pheretima elongata . . . Seminal vesicles. 

Stomatophora coronata . . . Pheretima barbadensis . Seminal vesicles. 

Stomatophora diadema . . . Pheretima barbadensis . Seminal vesicles. 

Pheretima posthuma . Seminal vesicles. 
Fam. Zygocystid^. 

Extremocystis dendrostomi. Dendrostoma signifer . CcElomic cavity. 


Aikinetocystis singularis . Eutyphoeus Joveatus . . Ccelomic cavity. 

Eutyphoeus spinulosus . Ccelomic cavity. 

Eutyphoeus rarus Ccelomic cavity. 

Eutyphoeus peguanus . . Ccelomic cavity. 

Nellocystis birmanica .... Pheretiyna compta Ccelomic cavity. 


Lankesteria culicis Aedes {Stegomyia) Stomach and Mal- 

segypti. pighian tubes. 

Aedes (Stegomyia) albo- Stomach and Mal- 

pictus. pighian tubes. 

Lankesteria mackiei Phlebotomus argentipes. Alimentary canal 

and hsemocoele. 

Phlebotomus papatasi . Alimentary canal 

and hsemocoele. 

Lankest-eria tripteroidesi . . Tripteroides dofleini., Hsemocoele and 

respiratory sys- 
Fam. Lecudestid^. 

Lecudina brasili Lumbriconereis sp. . . . Intestine. 

Bhatiella morphysse Morphysa sanguinea . Mid-gut. 

Ferraria cornucephali . . . Morphysa sanguinea . Mid-gut. 

Incert^ sedis. 

Dirhynchocystis globosa . . Eutyphoeus sp Seminal vesicles. 

Pheretima posthuma . . Seminal vesicles. 

Orayallia quadrispina. . . . Pheretima, heterochseta . Ccelomic cavity. 

* For Supplementary List of Parasites and their Hosts recorded since 
the above list was in type, see p. 374. 



Fam. Stenophorid^. 
Stenophora ellipsoidi .... 
Stenophora khagendras . . . 

Fam. Hyaxosporinid^. 
Hyalosporina cambolop- 

Hyalosporina rayi 

Fam. Gregarinid^. 
Leidyana gryllorum 



Diplopoda sp Mid-gut. 

Zikadesmus (?) sp Intestine. 

Gambolopsis sp Alimentary canal. 

Polydesmus sp Alimentary canal. 

Strongylosoma contor- Alimentary canal. 


Gryllus sp. 

Leidyana xylocopse . 
Gregarina aciculata 
Gregarina cornwalli. 
Gregarina oviceps . . 

Xylocopa sestuans . , 
Lepisnla saccharina 
Lepisma saccharina 
Gryllus sp 

Caulocephalus crenata .... 
Protomagalhsensia (?) 

Hirmocystis (?) parapene- 


Steinina metaplaxi 

Fam. Dactylophorid-j;. 
Grebneckiella nainllse .... 

Aulacophora foveicollis. 
Parapeneopsis sculptilis 

Gizzard and mid- 

Alimentary canal. 



Gizzard and mid- 

Alimentary canal. 


Parapeneopsis sculptilis Intestine. 

Metaplax dentipes 

Scolopendra sp 


Monoductus lunatus Strongylosoma contor- 



Fam. Adeleida. 
Adelea pachelebrse Pachelabra mcestra . . . 

Adelina schellacki Cormocephalus dentipes 


Heemogregarina berestneffi. 

Haen^ogregarina cantliei . . 
Haemogregarina hankini . 

Haemogregarina laverani . 

Haemogregarina leschen- 

Haemogregarina magna . . 

Haemogregarina mala- 

Haemogregarina mesnili . . 
Haemogregarina mirahilis . 

Rana hexadactyla . . . 
Eana limnocharis . . . 

Rana tigrina 

Eryx conicus 

Crocodilus porosus . . 
Oravialis gangeticu^ . 
Lissemys punctata 

Hemidactylus leschen- 

Rana limnocharis . . . 

Rana tigrina 

Lissemys punctata 


Kachuga tectum 

Tropidonotus asperri- 

Trovidonotus piscator 

Alimentary canal. 

Intestine and di- 
gestive glands. 




Blood, liver, bone- 
marrow, etc. 





Fam. H^MOGREGAnnsriD^ 

Hasmogregarina najse .... 
Hsemogregarina nicorise . . 

Hcsmogregarina nucleo- 

Hsemogregarina pythonis , 
Hsemogregarina rara .... 
Hsemogregarina rodriguesi. 
Hsejnogregarina stepanowi- 

Hsemogregarina testudinis. 
Hsemogregarina thomsoni . 
Hsemogregarina ihyrsoidea. 
Hsemogregarina tiedri . . . 
Hsemogregarina vittatse . . 

Hsemogergarina xavieri . . 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hsemogregarina sp 

Hepatozoon adiei 

Hepatozoon canis 



Hepatozoon felis domestici. 
Hepatozoon funambuli . . . 

Naja naja Blood. 

Geoemyda trijuga Blood. 

Lissemys punctata Blood. 


Ozobranchus shipleyi . Body. 

Bufo melanostictus . . . Blood. 

Python molurus Blood. 

Chinemys reevesii Blood. 

Heinidactylus brookei. . Blood. 

Chinemys reevesii Blood. 

Testudo emys Blood. 

Agama tuberculata . . . Blood. 

Thyrsoidea macrurus . Blood. 

Hemidactylus triedrus . Blood. 

Lissemys punctata Blood. 

Lissemys punctata Blood, lungs, 

granosa. spleen, liver. 

Trichogaster fasciatus . Blood. 

Varanus ^nonitor Blood. 

Bungarus cseruleus . . , Blood. 

Chrysopelea ornata .... Blood. 

Zaocys mucosus Blood. 

Porocephalus pattoni . . Body. 

Coluber helena Blood. 

Coluber sp Blood. 

Dipsadomorphus for- Blood. 

Dipsadomorphus ceylon- Blood. 

Dendrophys pictus . . . Blood. 

Dryophis mycterizans . Blood. 

Eryx johnii Blood. 

Naja bunga7'us Blood. 

Naja naja Blood. 

Naja naja var. atra . . . Blood. 

Naja sp Blood. 

Python sp Blood. 

Tropidonotus stolatus . . Blood. 

Vipera russellii Blood. 

Eagle (not identified) . Lung and leuco- 

Canis aureus Internal organs 

and leucocytes. 

Canis familiaris Internal organs 

and leucocytes. 

Cyon dukhunensis .... Internal organs 
and leucocytes. 

Vulpes bengalensis . . . Internal organs 
and leucocjrtes. 

Rhipicephalus san- Body. 

Felis sp Leucocytes. 

Funambulus pennantii. Leucocytes . 

Hsematopinus sp Body. 


Parasite. Host. Seat. 

Hepatozoon gerbilU Tatera indica Erythrocytes. 

Heematopinus stephensi Body. 

Hepatozoon leporis Lepus nigricollis Blood. 

Hepatozoon muris Rattus norvegicus .... Liver-cells and 


Rattus rufescens Liver-ceUs and 

Lselaps echidninus . . . Body. 

Hepatozoon sp Pteromys petaurista . . Blood. 

Karyolysus jorgei Calotes versicolor major Blood, and endo- 
thelial cells of 
liver and lungs. 
Fam. EiMERiiD^. 

Lankesterella minima .... Rana tigrina Blood. 

Lankesterella monilis .... Rana limnoclmris .... Blood. 

Rana tigrina Blood. 

Jsospora belli Homo sapiens Alimentary canal. 

Isospora bigemina Canis sp Sub-epithelial 

tissues of the 
Isospora caloti Calotes versicolor Intestinal epithe- 

Isospora felis Felis domesticus Intestinal epithe- 


Isospora knowlesi Hemidactylus fiaviviri- Alimentary canal, 


Isospora m,inuta Naja naja Alimentary canal. 

Isospora rivolta Felis domesticus Intestinal epithe- 

Herpestes auropunctaius Caecum. 

Isospora wenyoni Bufo melanostictus . . . Small intestine. 

Isospora sp Bos sp Alunentary canal. 

Eimeria clupearum Homo sapiens Intestine. 

Eimeria columbae Columba livia inter- Intestine. 


Eimeria cylindrica Natrix piscator Rectum. 

Eim,eria faurei Ovis sp Intestine. 

Eimeria flaviviridis Hemidactylus fiaviviri- Gall-bladder and 

dis. intestine. 

Eimeria harpodoni Harpodon nehereus . . . Alimentary canal. 

Eimeria hemidactyli Hemidactylus fiaviviri- Intestine. 

Eimeria kermorganti .... Gavialis gangeticus . . . Spleen. 

Eimeria knowlesi Hemidactylus fiaviviri- Intestine. 


Eimeria laminata Bujo melanostictus . . . Small intestine. 

Eimeria legeri Lissemys punctata Gall-bladder and 

granosa. bile-ducts. 

Eimeria m,itraria Chinemys reevesii Intestine. 

Eimeria najse Naja naja Rectum. 

Eimeria piscatori Natrix piscator Rectum. 

Eimeria sardinse Homo sapiens Alimentary canal. 

Eimeria southwelli ^tobatis narinari .... Intestine. 

Eimeria yakimovi Boselaphus tragoca- Alimentary canal. 


Eimeria sp Bos sp Alimentary canal. 

Eimeria sp Trichiurus savala .... Intestine. 

Eimeria sp Batrachus grunniens . . Intestine. 

Eimeria sp Epinephelus tauvina . . Intestine. 


Fam. EiMEBiiD^ (cont.). 

Eimeria sp 

Eimeria sp 

Eimeria sp 

Eimeria sp 

Eimeria sp , 

Eimeria sp 

Eimeria (?) sp 

Wenyonella hoarei 

Pythonella bengalensis. , . . 
Aggregata sp 

Incert^ sedis. 

Toxoplasma canis 

Toxoplasma columbae . . . . 
Toxoplasma cuniculi . . . . 

Toxoplasma fulicie 

Toxoplasma sp 

Toxoplasma sp 

Toxoplasma (1) pyrogenes . 


Hsemoproteus antigone . . . 
Hmmoproteus asturis . . . . 

Hsemoproteus cerchneisi . . 

Hsemoproteus columbae . . . 

Hsemoproteus coracise . . . 

Hsemoproteus corvusi .... 

Hsemoproteus danilewskyi 




Engraulis mystax .... Intestine. 

Otolithus ruber Intestine. 

Sillago sihama Intestine. 

Coilia dussumieri Intestine. 

Plotossus canius Intestine. 

Epinephelus diacanthv^ Intestine. 

Culex sp Stomach cells. 

Sciurus sp Intestine. 

Python sp Intestine. 

Parapeneopsis sculp- Intestine. 

Canis familiaris Liver and spleen. 

Columba livia Leucocytes. 

Lepus sp Liver, spleen,bone- 

marrow, and 

Fulica atra Lung. 

Passer sp Blood. 

Saxicola caprata Blood. 

Homo sapiens Blood and spleen. 

Hsemoproteus glaucidii . . 
Hsemoptoteus gymnorhidis. 
Hsemoproteus herodiadis . 

Hsemoproteus kopki 

Anthropoides virgo , . . 
Astur badius dussu- 
Cerchneis tinnunculo- 

sus objurgatus. 

Columba livia 

Lynchia maura 

Coracias benghalensts 

Corvus levaillanti 

macrorhynchus . 

Corvus splendens 

Anas {Fuligula) baeri . 
Copsychus saularis . . . 
Coracias benghalensis 

Garrulax albigularis . . 
Garrulus lanceolatus . . 
Glareola pratincola .... 
Kittacincla macroura . 
Melophus melanicterus 
Mesia argentauris .... 
Nettapus coromandeli- 

Otus bakkamcena var. 

Propasser rhodochrous . 
Prunella strophiata jer- 

Glaucidium radiatum . 
Oymnoris xanthocoUis . 
Egretta intermedia 

Hemidactylus brooki . . 
















Haemoproteus machlolophi. 


Machlolophus xantho- 



Hsemoproteus metchnikovi. 

Chitra indica 


Hxmoproteus orioli ..... 

Or-iolus oriolus kundoo . 


Hsemoproteus raymundi . 

Leptocoma zeylonica . . 

Blood, lungs, 

Hsemoproteua rileyi 

Pavo cristatus 


Haemoproteus simondi . . . 

Hemidactylus leschen- 


Hsemoproteus sturni 

Sturnia malaharica . . . 


Hsemoproteus upupse .... 

Upupa epops orientalis. 

Blood, lungs, 

Hsemoproteus wenyoni . . . 

Orthotomus sutorius . . 


Hsemoproteus sp. ... 

4n^^g^o?^e antigone .... 


Hsemoproteus sp 

Calcenas nicobarica . . . 


Hsemoproteus sp 

Centropus sinensis. , . . 


Hsemoproteus sp 

Chloropsis aurifrons . . 


Hsemoproteus sp 

Turdoides striatus 


Haemoproteus sp 

Cyanops flavifrons . . . 


Hsem,oproteus sp 

Dendrocitta rufa vaga- 


Haemoproteus sp 

Haliaeetus leucoryphus. 


Hsemoproteus sp 

Hemidactylus sp 


Hsemoproteus sp 

Coryllis beryllinus .... 


Hsemoproteus sp 

Psittacula alexandri . . 


Haemoproteus sp 

Otus bakkamoena 


Haemoproteus sp 

Strix ocellata 


Hsemoproteus (?) segithinse. 

Mgithina tiphia 


Hsemoproteus (?) anthi . . . 

Anthus richardi rufulus 


Haemoproteus (?) hranse . . 

Athena brama 


Haemoproteus (?) centropi . 

Centropus sinensis 


Haemoproteus (?) dicruri. . 

Dicrurus macrocercus 


Haemoproteus (?) efew . . . 

Elanus cseruleus voci- 


Haemoproteus (?) galUnulae 

Gallinula chloropus . . . 


Haemoproteus (?) halcyonis 

Halcyon smyrnensis. . . 


Hsemoproteus (?) otocompsae 

Elathea jocosa 


Hsemoproteus (?) pastoris . 

Pastor roseus 


Haemoproteus (?) platalese. 

Platalea leucorodia 


Hsemoproteus (?) tephro- 

Tephrodornis pondiceri- 



anus pondicerianus. 

Hsemoproteus (?) therei- 

Thereiceryx zeylanicus 




Haemoproteus (?) therei- 

Thereiceryx zeylanicus 


cerycis var. zeylonica. 


Leucocytozoon chloropsidis. 

Chloropsis aurifrons 


Leucocytozoon coraciae. . . . 

Coracias benghalensis 

Blood and smears 


from limgs. 

Leticocytozoon melloi .... 

Coracias benghalensis 

Blood and smears 


from lungs. 

Leucocytozoon sp 

Athene brama 


Leucocytozoon sp 

Leucocytozoon sp 

Falco sp 


lanthocincla rufogularis 


Leucocytozoon sp 

Leiothrix lutea 




Parasite. Host. 

Fam. H^MOPROTEiD^ (cont.). 

Leucocytozoon sp Oreicola ferrea 

Leucocytozoon sp Propasser rhodochrous 

Leucocytozoon sp Peking robins 


Chloropsis aurifrons 


Coluinba sp 

Gallinula chloropus . 
Egretta intermedia • 

ter media. 
Gopsychus saularis . 

Fam. Plasmodid^. 
JProteosoma chloropsidis 

Proteosoma columbse . . 
Proteosoma gallinulse . . 
Proteosoma herodiadis. . 

Proteosoma moruony . . 

Proteosoma passeris Passer domesticus .... 

Proteosoma prsecox Aidemosyne malaharica 

Sturnus vulgarus pol- 

Tragopan satyra 

Anopheles suhpictus . . 

Culex fatigans 

Proteosoma sip. Antigone antigone .... 

Proteosoma sp Corvus splendens 

Proteosoma sp Coryllis beryllinus .... 

Proteosoma sp Cyanops flavifrons . . . 

Proteosoma sp Elathea jocosa 

Proteosoma sp Emberiza fucata 

Proteosoma sp Gracula indica 

Proteosoma sp Garrulax leucolophus . 

Proteosoma sp Leiothrix lutea 

Proteosoma sp Melophus melanicterus 

Proteosoma sp Saxicola caprata 

Proteosoma sp Turdus boulboul 

Lavei'ania malarise Homo sapiens 

Anopheles annularis . . 

Anopheles culicifacies . 

Anopheles fluviatilis . 

Anopheles funestus . . . 

Anopheles jeyporiensis . 

Anopheles maculatus . . 

Anopheles minimus . . 

Anopheles philippinen- 

Anopheles stephensi . . 

Anopheles sundaicus . . 
Plasmodium vivax Homo sapiens 

Anopheles annularis . . 

Anopheles culicifacies . 

Anopheles fluviatilis . . 

Anopheles funestus . . . 

Anopheles Jeyporiensis . 

Anopheles maculatus . 

Anopheles minimus . . 

Anopheles philippinen- 

Anopheles stephensi . 

Anopheles sundaicus . 

Homo sapiens 

Anopheles annularis . 

Anopheles culicifacies 

Plasmodium malarise 











ium malariae . . . 

Anopheles fluviatilis . . 
Anopheles funestus . . . 
Anopheles jeyporiensis. 
Anopheles maculatus . 
Anopheles minim,us . . 
Anopheles philippinen- 




Anopheles stephensi . . Body. 

Anopheles sundaicus . . Body. 

Plasmodium cynomolgi . . Silenus irus Blood. 

Silenus rheSus Blood. 

Silenus sinicus Blood. 

Anopheles annularis . . Salivary glands. 

Anopheles culicifacies . Salivary glands. 

Anopheles maculatus . Salivary glands. 

Anopheles splendidus . Salivary glands. 

Plasmodium inui Silenus irus Blood. 

Silenus rhesus Blood. 

Plasmodium knowlesi . . . Silenus irus Blood. 

Silenus rhesus Blood. 

Plasmodium kochi ...... Cercopithecus spp Blood. 

Plasmodium pithed Simia satyrus Blood. 

Plasmodium semnopithecus Presbytes pileatus .... Blood. 

Pygathrix entellus .... Blood. 

Silenus rhesus Blood. 

Plasmodium bubalis Bos indicus Blood. 

Bubalus sp Blood. 

Plasmodium canis Canis familiaris Blood. 

Plasmodium equi Equus caballus Blood. 

Plasmodium mackiei .... Myotis muricola Blood. 

Plasmodium narayani . . . Lutra lutra Blood. 

Plasmodium pteropi Pteropus m,edius Blood. 

Plasmodium ratufee Ratiifa indica Blood. 

Plasmodium tyrio Manes pentadactyla . . Blood. 

Fam. Theileriid^. 

Theileria celli Macacus pileatus .... Blood. 

Theileria hirci Capra hircus Blood. 

Theileria mutans Bos indicus Blood. 

Theileria parva Bos indicus Blood. 

Fam. Babesiid^. 

Babesia bigemina Bos indicus Blood. 

Babesia caballi Equus caballus Blood. 

Babesia canis Canis aureus Blood. 

Canis familiaris Blood. 

Rhipicephalus sangui- Blood. 

Babesia equi Equus caballus Blood. 

Babesia felis Felis sp Blood. 

Babesia gibsoni Canis aureus Blood. 

Canis familiaris Blood. 

Babesia motasi Ovis sp Blood. 

Babesia ninense Erinaceus sp Blood. 

Babesia sergenti Capra hircus Blood. 

Babesia soricis Crocidura ceerulea .... Blood. 

Babesia tropicus Elephant, camel, gui- Blood. 

nea-pigs, fowls, and 


Fam. Babesiid^ (conL). 

Babesia sp 

Babesia sp 

Babesia sp 

Incert^ sedis. 

Grahamella muris 

Grahamella sp 

Paraplasma sp 

Anaplasma sp 



Axis axis 

Cyon dukhunensis . 
Herpestes edwardii 


Manes pentadactyla . 

Homo sapiens 


Chitra indica 

Coluber blumenbachii. 
Hemidactylus brooki . 

Homo sapiens 

Naja naja 

Perca fluviatilis 

Rana esculenta 

Rana tigrina 





Fam. Ceratomyxid^. 

Ceratomyxa gobioidesi. . . 

Chloromyxum amphinovi 
Fam. Sph^rosporid^. 

Sphasrospora sp 

Fam. Myxidiid^. 
Myxidium, danilewshyi . 

Myxidium mackiei 

Myxidium sp 

Zschokkella prashadi , 

Gobioides rubicundus . Liver, gall-bladder 
kidney, ovary. 

Macrones gulio Liver,gall-bladder, 

kidney, ovary. 

Trichogaster fasciatus 
Am,phipnous kuchia . 


kidney, ovary. 

Barilius barna Under the scales. 

Fam. Myxobolid^. 
Myxobolus calbasui 

Myxobolus nodularis. . . . 
Myxobolus sp 

Thelohanellus rohitas . . . 
Thelohanellus seni 

Henneguya ophiocephali. 

Chinemys reevesii. . 
Trionyx gangeticus . 
Clarias batrachus . 
Geoemyda trijuga . 
Kachuga smithi . . . 
Lissemys punctata 

Ophiocephalus punctatus 
Saccobranchus fossilis . 
Bufo m,elanostictus . . . 
Lissemys punctata 

Rana tigrina 

Cirrhina mrigala 

Clarias batrachus . 

Katla katla 

Rasbora daniconius 
Rasbora daniconius 

Labeo rohita 
Labeo rohita 

Ophiocephalus puncta- 









Liver and gall- 

Ovary and liver. 



Subcutaneous tis- 


Median and caudal 

Gills and muscles. 




Henneguya otolithus 

Host. Seat 

Otolithus maculatus . . . Bulbus arteriosus. 
Otolithus ruber Bulbus arteriosus. 


Fam. NosEMATiD^. 
Nosema adiei 

Cimex rotundatus 

Nosema bombycis. . . 
Nosema ctenocephali 

Thelohania indica , 

Thelohania legeri . . 

Thelohania obscura 

Microsporidian {indet.) 

Microsporidian (indet.) 

Fam. Sakcocystid^ 
Sarcocystis blanchardi . . 

Sarcocystis lindemanni 

Fam. Globidid^. 
Globidium, fuisiformis . . 
Globidium sp 


Incert^ sedis. 
Rhinosporidium seeberi, 
Ehinosporidium sp 

Bombyx mori 

Ctenocephalus canis . . 

Anopheles hyracanus . 

Anopheles barbirostris . 

Anopheles fuliginosus , 

Anopheles funestus . . . 

Anopheles hyracanus . 

Anopheles pseudojamesi 

Anopheles ramsayi .... 

Anopheles subpictus . . . 

Anopheles funestus , . . 

Stegomyia sp 

Culex fatigans 

Gut, salivary 

glands, and 

All tissues. 
Digestive tract of 

Adipose tissue of 

Adipose tissue of 

Adipose tissue of 

Adipose tissue of 

Adipose tissue of 

Adipose tissue of 

Adipose tissue of 

Adipose tissue of 

Adipose tissue of 

Nerve -cord of 

Nerve-cord of 


Bos bubalus Muscles. 

Bos indicus Heart -muscle. 

Homo sapiens Muscles. 

Bos indicus Alimentary canal. 

WaUaby Alimentary canal. 

Homo sapiens Nasal polypi. 

Homo sapiens Abscess cavities. 

(ii) List of Hosts and their Parasites *. 
Host. Parasite. Seat. 


Axis axis Babesia sp Blood. 

Bat Anaplasma sp Blood. 

Bos bubalus Plasmodium bubalis. . . Blood. 

Sarcocystis blanchardi . Muscles. 

* For Supplementary List of Hosts and their Parasites recorded sine© 
the above list was in type, see p. 375. 


Host. Parasite. Seat. 

Mammalia (cont.). 

Bos indicus Plasmodium bubalis. . . Blood. 

Theileria mutans Blood. 

Theileria parva Blood. 

Babesia bigemina Blood. 

Sarcocystis blanchardi . Heart-muscle. 

Globidium Jusiformis . Alimentary canal. 

Bos sp Isospora sp Alimentary canal. 

Eimeria sp Alimentary canal. 

Camelus sp Babesia tropicus .... Blood. 

Canis aureus Hepatozoon canis .... Internal organs 

and leucocytes. 

Babesia canis Blood. 

Babesia gibsoni Blood 

Canis familiaris Hepatozoon canis .... Internal organs 

and leucocytes. 

Toxoplasma canis . . . Blood. 

Plasm^odium canis .... Blood. 

Babesia canis Blood. 

Babesia gibsoni Blood. 

Canis sp Isospora begemina .... Subepithelial tis- 
sues of the in- 

Capra hircus Theileria hirci Blood. 

Babesia sergenti Blood 

Cavia porcellu^ (?) Babesia tropicus Blood. 

Cercopithecus sp Plasmodium kochi .... Blood. 

Crocidura cserulea Babesia soricis Blood. 

Cyon dukhensis Hepatozoon canis .... Internal organs 

and leucocytes. 

Babesia sp Blood. 

Elephas sp Babesia tropicus Blood. 

Equus caballus Plasmodium equi .... Blood. 

Babesia caballi Blood. 

Babesia equi Blood. 

Erinaceus sp Babesia ninense Blood. 

Felis domesticus Isospora felis Intestinal epithe- 

Isospora rivolta Intestinal epithe- 


Felis sp Hepatozoon felis Leucocytes. 


Babesia felis Blood. 

Funambulus pennantii .. Hepatozoon funanibuli. Leucocytes. 

Herpestes auropunctatus . . Isospora rivolta Caecum. 

Herpestes edwardii Babesia sp Blood. 

Homo sapiens Isospora belli Alimentary canal. 

Eimeria clupearum . . . Intestine. 

Eimeria sardinss Ahmentary canal. 

Toxoplasma (?) pyro- Blood and spleen. 

Laverania malariee . . Blood. 

Plasmodium malarise. . Blood. 

Plasmodium vivax . . . Blood. 

Paraplasma sp Blood. 

Anaplasma sp Blood. 

Sarcocystis lindemanni. Blood. 

Rhinosporidium seeberi Nasal polypi. 

Rhinosporidium sp. . . Abcess cavities. 



Lepus sp. . 



Lepus nigricollis . . . 

Lutra Intra 

Macacus pileatus . . . 
Manes pentadactyla 

Myotis muricola 
Ovis sp 

Presbytes pileatus . . 

Pteromys petaurista . 
Pteropus medius . . . 
Pygathrix entellus . . 

Raftus norvegicus. 
Rattus rufescens . 

Rattus sp. . . , 
Ratufa indica 
Sciurus sp. 
Silenus irus . 

Silenus rhesus 

Silenus sinicus . . . 
Simia satyrus . . . . 
Tateria indica . . . . 
Vulpes bengalensis 



Mgithina tiphia 

Aidemosyne malabarica 
Anas (Fuligula) baeri . . 

Anthropoides virgo . . . . 
Anthus richardi rufulus 
Antigone antigone . . . . 

Astur badius dussumieri 
Athene brama 

Calcenas nicobarica 

Centropus sinensis parroti 
Cerchneis tinnunculosus 

Chloropsis aurijrons .... 

Toxoplasma cuniculi . . 

Liver, spleen, 


and heart-blood, 

Hepatozoon leporis . . . 


Plasmodium narayani 


Theileria celli 


Plasmodium tyrio 


Chuhamella sp 


Plasmodium maclciei . . 


Eimeria fatirei 


Babesia motasi 


Plasmodium se7nnopi- 



Hepatozoon sp 


Plasmodimn pteropi . . . 


Plasmodium semnopi- 



Hepatozoon muris .... 

Liver-cells and 


Hepatozoon muris .... 

Liver-cells and 


Grahamella muris 


Plasmodium ratufse . . 


Wenyonella hoarei . . . 


Plasmodium cynomolgi 


Plasmodium inui .... 


Plasmodium knowlesi . 


Plasmodium cynomolgi. 


Plasmodium inui .... 


Plasmodium knowlesi . 


Plasjnodiuin semnopi- 



Plasmodium cynomolgi. 


Plastnodium pitheci . . 


Hepatozoon gerbilli . . . 


Hepatozoon canis .... 

Internal organs 

and leucocytes. 

Olobidium sp 

Alimentary canal. 

Hsemopr-oteus (?) segi- 



Proteosoma prsecox .... 


Hssmoproteus danilew- 



Heemoproteus antigone . 


Hseinoproteus (?) anthi. 


H£e7no]}roteiis sp 


Proteosoma sp 


Heemoproteus asturi. . . 


Hsemoproteus (?) b7'am£e 


Leucocytozoon sp 


Hsemopi'oteus sp 


Hsemoproteus (?) sp. . . 


Hsemoproteus ceixhneisi 


Hsemoproteus sp. 




AvES (cont.). 
Chloropsis aurifrons david- 

Columba livia 

Columba livia intermedia . 

Columba sp 

Copsychiis saularis 

Coracias benghalensis ben- 

Coracias benghalensis in- 

Corvus levaillanti macro- 

Corvus splendens 

Coryllis beryllinus 

Cyanops flavijrons 

Dendrocitta rufa vagabunda 
Dicrurus macrocercus macro- 

Egretta intermedia inter- 
Elanus cseruleus vociferus . 
Elathea jocosa 

Emberiza fucata 

Falco sp 

Fulica atra 

Gallinula chloropus 

Callus gallus 

Garrulax albigularis 

Oarrulax leucolophus .... 
Garrulus lanceolatus .... 

Qlareola pratincola 

Olaucidium radiatum .... 

Gracula indica 

Gymnoris xanthocollis . . . 

Halcyon smyrnensis 

Haliaeetus leucoryphus . , . 




Leucocytozoon chlorop- 



Proteosoma chloropsidis. 


Toxoplasma columbee . 


Hsemoproteus columbee . 


Eimeria columbse .... 


Proteosoma columbse . . 


Heemoproteus danilew- 



Proteosoma moruony . . 


Heemoproteus coracise . 


Leucocytozoon coracise . 

Blood and smears 

from lungs. 

Leucocytozoon melloi . . 

Blood and smears 

from lungs. 

Hsemoproteus sp 


Heemoproteus corvusi . . 


Heemoproteus corvusi . 


Proteosoma sp 


Hsemoproteus sp 


Proteosoma sp 


Hsemoproteus sp 


Proteosoma sp 


Hsemoproteus sp 


Hsemoproteus (?) dicruri 


Hepatozoon adiei 

Ltmgs and leuco- 


Heemoproteus herodiadis 


Proteosoma herodiadis . 


Heemoproteus (?) elani. 


Hsemoproteus (?) oto- 



Proteosoma sp 


Proteosoma sp 


Leucocytozoon sp 


Toxoplasma fulicse . . . 


Heemoproteus (?) galli- 



Proteosoma gallinulse . 


Babesia tropicus 


Hxmoproteus danilew- 



Proteosoma sp 


Hsemoproteus danilew- 



Hsemoproteus danilew- 



Heemoproteus glaucidii. 


Proteosoma sp 


Heemoproteus gymno- 



Hssynoproteus (?) halcy- 



Hsemoproteus sp 




Host. Parasite. 

lanthocincla rufogularis . Leucocytozoon sp 

Kittacincla macroura .... Hsemoproteus danilew- 

Leptocoma zeylonica Hsemoproteus raymundi 

Leiothrix luteus Leucocytozoon sp 

Proteosoma sp 

Maclolophus xanthogenys . Hsemoproteus machlo- 

Melophtis melanicterus . . Hsemoproteus danilew- 


Proteosoma sp 

Mesia argentauris Hsemoproteus danilew- 

Nettapus coromandelianus. Hsemoproteus danilew- 


Oreicola ferrea Leucocytozoon sp 

Oriolus oriolus kundoo . . Hsemoproteus orioli . . 

Orthotomus sutorius Hsemoproteus wenyoni . 

Otus bakkamoena Hsemoproteus sp 

Otus bakkamoena var. Hsemoproteus danilew- 

malabarica. skyi. 

Passer domesticus Proteosoma passeris . . 

Passer sp Toxoplasma sp 

Pastor roseus Hsemoproteus (?) pas- 


Pavo cristatus Hsemoproteus rileyi . . 

Peking robins Leucocytozoon sp'. .... 

Platalea leucorodia major. Hsemoproteus (?) plata- 

Propasser rhodochrozis . . . Hsemoproteus danilew- 

Prunella strophiatajerdoni Hsemoproteus danilew- 


Psittacula alexandri fasci- Hsemoproteus sp 


Leucocytozoon sp 

Saxicola caprata ........ Toxoplasma sp 

Proteosoma sp 

Strix ocellata Hsemoproteus sp 

Sturnia malabarica ..... Hsemoproteus sturni . . 
Sturnus vulgarus poltarat- Proteosoma prsecox .... 

Tephrodornis pondicerianus Hsemoproteus ( ? ) tephro - 

pondicerianus. dornis. 

Thereiceryx zeylanicus in- Hsemoproteus (?) therei- 

ornata. cerycis. 

Thereiceryx zeylanicus zey- Hsemoproteus (?) therei- 
lanicus. cerycis var. zeylonica. 

Tragopan satyra Proteosom,a prseeosx . . 

Turdoides striatus striatus . Hsemoproteus sp 

Turdus boulboul Proteosoma sp 

Upupa epops orientalis . . Hsemoproteus upupse . 


Agama tvberculata Hsemogregarina thom- 

Bungarus coeruleus Hsemogregarina sp. ... 


Blood, lungs, 







Blood, lungs, liver. 















Reptilia (cont.). 
Calotes versicolor . . . 




Calotes versicolor major 
Chinemys reevesii 

Isospora caloti 

Karyolysiis jorgei .... 

Ghitra indica 

Chrysopelea ornata 

Coluber hlumenbachii .... 

Coluber helena 

Coluber sp 

Crocodilus porosus 

Dendrophys pictus 

Dipsadomorphus ceylonen- 

Dipsadomorphus forstenii . 
Dryophis mycterizans .... 

Eryx conicus 

Eryx johnii 

Gavialis gangeticus 

Geoemyda trijuga 

Hemidactylus brooki .... 

Hemidactylus flaviviridis 

Hemidactylus leschenaulti . 

Hemidactylus triedrus . . . 

Hemidactylus sp 

Kachuga smithi 

Kachuga tectum 

Lissemys punctata granosa. 

Intestinal epithe- 
Blood, and endo- 
thelial cells of 
liver and lungs. 

Hsemogregarina rara . . Blood. 

Hsemogregarina stepa- Blood. 

Eimeria mitraria Intestine. 

Myxidium danilewskyi. Kidney. 

Hsemoproteus metchni- Blood. 

Anaplasma sp Blood. 

Hsemogregarina sp. . . . Blood. 

Anaplasma sp Blood. 

Hsemogregarina sp. . . . Blood. 

Hsemogregarina sp. . . . Blood. 

Hsemogregarina hankini Blood. 

Hsemogregarina sp. . . . Blood. 

Hsemogregarina sp. . . . Blood. 

Hsemogregarina sp. . . . Blood. 

Hse'inogregarina sp. . . . Blood. 

Hsemogregarina cantliei Blood. 

Hsemogregarina sp. . . . Blood. 

Hsemogregarina han- Blood. 


Ei?neria kermorganti . . Spleen. 

Hsemogregarina nicorise Blood. 

Myxidium sp GaU-bladder. 

Hsemogregarina rodri- Blood. 


Hsetnoproteus kopki . . Blood. 

Anaplasma sp Blood. 

Isospora knowlesi .... Alimentary canal. 

Eimeria flaviviridis . . . Gall-bladder and 


Eitneria hemidactyli. . . Intestine. 

Eimeria knowlesi Intestine. 

Hsemogregarina leschen- Blood. 


Hsemoproteus simondi . Blood. 

Hsemogregarina tiedri . Blood. 

Hsemoproteus sp Blood. 

Isospora knowlesi Gall-bladder. 

Hxmogregarina mesnili Blood. 

Hsemogregarina laver- Blood. 


Hsemogregarina mala- Blood, liver, bone- 

barica. marrow, etc. 

Hsemogregarina nicorise Blood. 

Hsemogregarina vittatse. Blood. 

Hsemogregarina xavieri. Blood, lungs, 
spleen, liver. 

Eimeria legeri Gall-bladder and 


Myxidium sp Gall-bladder. 

Zschokkella prashadi . Gall-bladder. 







Babesia tropicus 


Naja bungarus 

Hmmogregarina sp. ... 


Naja naja 

Hsemogregarina najse . 


Haemogregarina sp. ... 


Isopsora minuta 

Alimentary canal. 

Eimeria najse 


Anaplasma sp 


Naja naja var. atra 

Hsemogregarina sp. ... 


^070 sp 

Heemogregarina sp. ... 


Natrix piscator 

Eimeria cylindrica . . . 


Eimeria piscatori .... 


Python molurus 

Hsemogregarina pytho- 


Python sp 

Hsemogregarina sp. ... 


Pythonella bengalensis . 


Testudo emys 

Hsemogregarina testu- 


Trionyx gangeticus 

Myxidium machiei . . . 


Trop-idonotus asperrimus . 

Hsemogregarina mira- 


Tropidonotus piscator . . . 

Hsemogregarina mira- 


Tropidonotus stolatus . . . . 

Hsemogregarina sp. ... 


Varanus monitor 

Hsemogregarina sp. ... 


Vipera rtcssellii 

Hsemogergarina sp. ... 


Zaocys mucosus 

Hsemogregarina sp. ... 



Bufo melanosticttis 

Hsemogregarina nucleo- 


Isospora wenyoni .... 

Small intestine^ 

Eimeria laminata .... 

Small intestine- 

Zschokkella prashadi . 


Rana e-sculenta 

Anaplasma sp 


Rana hexadactyla 

Hsemogregarina berest- 


Rana limnocharis 

Hsemogregarina berest- 


Hsemogregarina magna. 


Lankesterella monilata . 


Rana tigrina 

Hsemogregarina berest- 


Hsemogregarina magna. 


Lankesterella minima . 


Lankesterella inonilata . 


Anaplasma sp 


Zschokkella prashadi . 



^tobatis narinari 

Eimeria southwelli . . . 


Amphipnaus kiichia 

Chloromyxum amphi- 


Bariliits barna 

Sphserospora sp 

Under the scales.- 

Batrachus grunniens .... 
Cirrhina mrigala 

Eimeria sp 


Myxobolus calbasut . . . 


Clarias batrachus 

Myxobolus calbasui . . . 

Ovary and liver. 

Myxidium sp 


Goilia diissumieri 

Eimeria sp 






Pisces {cont.). 

Engraulis mystax 

Epinephelus diacanthus . 
Epinephelus tauvina . . . 
Gobioides rubicundus . . . 

Harpodon nehereus . . . . 

Katla katla 

Labeo rohita 

Macrones gulio 

Ophiocephalus punctatus 

Otolithus ruber 

Plotossiis canius 

Rasbora daniconius . . . . 

Saccobranchus fossilis . . . 

Sillago sihama 

Thyrsoidea macrurus . . . 

Trichiurus savala 

Trichogaster fasciattis . . . 



Eiraeria sp Intestine. 

Eimeria sp Intestine. 

Eimeria sp Intestine. 

Geratomyxa gobioidesi . Liver, gall-bladder 
kidney, ovary. 

Eimeria harpodoni . . . Alimentary canal. 

Myxobolus sp Gills. 

Thelohanellus rohitse . . GUIs. 

Thellohanellus seni .... Median and caudal 

Geratomyxa gobioidesi . Liver, gaU-bladder 
kidney, ovary. 

Henneguya sp Gills and muscles. 

Myxidium sp GaU-bladder. 

Eimeria sp Intestine. 

Henneguya otolithus . . Bulbus arteriosus. 

Eimeria sp Intestine. 

Myxobolus nodularis . Muscles. 

Myxobolus sp Subcutaneus tis- 

Myxidium- sp GaU-bladder. 

Eimeria sp Intestine. 

Heemogregarina thyrsoi- Blood. 

Eimeria sp Intestine 

Heemogregarina sp. . . . Blood. 

Geratomyxa gobioidesi . Liver, gaU-bladder 
kidney, ovary. 


Pachelabra mcBstra 

Adelea pachelebrse .... 

Intestine and di- 
gestive glands. 


Lselaps echidninus 

Hepatozoon muris .... 


Porocephalus pattoni .... 

Heemogregarina sp. ... 


Rhipicephalus sanguineus . 

Hepatozoon canis .... 



Aedes (Stegomyia) segypti. 

Lankesteria culicis . . . 

Stomach and Mal- 
pighian tubes. 

Laverania malarise . . . 


Aedes (Stegomyia) albopic- 

Lankesteria culicis . . . 

Stomach and Mal- 


pighian tubes. 

Anopheles annularis .... 

Plasmodium cynomolgi. 

Salivary glands. 

Plasmodium malarise. . 


Plasmodium vivax .... 


Anopheles barbirostris . . . . 

Thelohania legeri 

Adipose tissue of 

Anopheles culicifacies. . . . 

Laverania malarias . . . 


Plasmodium vivax . . . 


Plasmodium malarise . . 


Plasmodium cynomolgi. 

Salivary glands. 

Anopheles fluviatilis . . . . 

Laverania malarise . . . 


Plasmodium vivax . . . 


Plasmodium malarise . . 


Anopheles fuliginosus . . . 

Thelohania legeri 

Adipose tissue of 



Anopheles funestus 

Laverania malarise . 
Plasmodium malarise 
Plasmodium, vivax . 
Thelohania legeri . . . 

Anopheles hyracanus . . 

Anopheles jeyporiensis 
Anopheles m^culatus . . 

Anopheles m,inimus 

Anopheles philippinensis 

Anopheles pseudojam,esi 
Anopheles ramsayi 

Thelohania obscura . 
Thelohania indica . . 
Thelohania legeri . . . 

Laverania malarias . 
Plasmodium malarise 
Plasmodium vivax . 
Laverania malarise . 
Plasmodium, malarise 
Plasmodiutn vivax . 
Plasmodium cynomolgi 
Laverania malaria . 
Plasmodium malarise 
Plasmodium, vivax . 
Laverania malarise . 
Plasmodium malarise 
Plasmodium vivax . 
Thelohania legeri . . . 

Thelohania legeri 

Anopheles splendidus . . . 
Anopheles stephensi .... 

Anopheles sundaicus 
Anopheles subpictus . 

Plasmodium cynomolgi. 
Laverania malaria 
Plasmodium malarise . 
Plasmodium vivax . 
Laverania malarise . 
Plastnodium malarise 
Plasmodium vivax . 
Proteosoma prxcox . . 
Thelohania legeri . 

Aulacophora foveicollis 

Bombyx mori 

Cimex rotundattis 

Gaulocephalus crenata 
Nosenia bombycis . . . 
Nosema adiei 

Ctenocephalus canis Nosema ctenocephali 

Proteosoma prsecox .... 
Microsporidian (indet.) 

Culex sp. . 
Gryllus sp. 

Hsematopinus stephensi 

Hsematopinus sp 

Lepisma saccharina . . . 

Phlebotomus argentipes 

Eimeria (?) sp 

Leidyana gryllorum . . 

Gregarina oviceps . . . . 

Hepatozoon gerbilli . . . 
Hepatozoon funambuli . 
Qregarina aciculata . . , 
Gregarina cormvalli . . 
Lankesteria niackiei . . 

Adipose tissue of 

Adipose tissue of 

Adipose tissue of 

Adipose tissue of 


Salivary glands. 
Adipose tissue of 

Adipose tissue of 

Salivary glands. 
Adipose tissue of 

Alimentary canal. 
All tissues. 
Gut, salivary 

glands, and 

Digestive tract of 

Nerve cord of 

Stomach cells. 
Gizzard and mid- 
Gizzard and mid- 
Alimentary canal 

and haemoccele. 



Host. Parasite. Seat. 

INSECTA {cont.). 

Phlebotomus papatasi . . . Lankesteria mackiei . . Alimentary canal 

and hasmoccele. 

Stegomyia sp Microsporidian {indet.) Nerve cord of 


Triptroidesi dofleini Lankesteria tripterodesi. Haemoccele and 

respiratory sys- 

Xylocopa sestuans lAedyana xylocopse . . . Gizzard and mid- 

Combolopsis sp. Hyalosporina combo- Alimentary canal. 


Cormocephalus dentipes . . Adelina schellacki .... Intestine. 

Diplopoda sp Stenophora ellipsoidi . Mid-gut. 

Polydesmus sp Hyalosporina rayi . . . Alimentary canal. 

Scolopendra sp Grebneckiella navillse . Intestine. 

Strongylosoma contortipes . Monoductus lunatiis .. Alimentary canal. 

Hyalosporina rayi . . . Alimentary canaL 

Zikadesmus (?) sp Stenophora khagendrse. Mid-gut. 


Metaplax dentipes Steinina metaplaxi . . . Intestine. 

Parapeneopsis sculptilis . Protomagalhaensiu Intestine. 

(?) attenuata. 

Hirmocystis (?) para- Intestine. 

Aggregata sp Intestine. 


Lumbriconereis sp Lecudina brasili Intestine. 

Morphysa sanguinea Bhatiella tnorphysse .. Mid-gut. 

Ferraria cornucephali . Mid -gut. 


Allolobophora caliginosa . . Ehynchocystis cognetti . Seminal vesicles.. 

Allolobophora fostida .... Nematocystis pluri- Seminal vesicles.. 


Eutyphoeus foveatus Aikinetocystis singu- Coelomic cavity. 


Eutyphoeus incommodus . . Nematocystis Stephen- Seminal vesicles.. 


Eutyphoeus nicholsoni . . . Monocystis beddardi . . Seminal vesicles.. 

EutyphcBus peguanus .... Aikinetocystis singu- Coelomic cavity. 


Eutyphoeus rarus Aikinetocystis singu- Coelomic cavity. 


Eutyphoeus spinulosus . . . Aikinetocystis singu- Coelomic cavity. 


Eutyphoeus sp. Monocystis beddardi . . Seminal vesicles. 

Dirhynchocystis globosa. Seminal vesicles.. 

Megascolex trilobatus .... Apolocystis mattliai . . . Seminal vesicles. 

Pheretima barbadensis . . . Nematocystis vermicu- Seminal vesicles. 

Stomatophora coronata. Seminal vesicles. 
Stomatophora diadema. Seminal vesicles.. 
Pheretima comptu Nellocystis birmanica . Coelomic cavity. 


Host. Parasite. Seat. 

Pheretima elongata Rhynchocystis awatii . . Seminal vesicles. 

Rhynchocystis mamil- Seminal vesicles. 


Stomatophora bulbifera. Seminal vesicles. 

Pheretima heterochseta . . . Nematocystis hessei . . . Seminal vesicles. 

Nematocystis lumbri- Seminal vesicles. 


Grayallia quadrispina . Ccelomic cavity. 

Pheretima posthuma Monocystis bengalensis. Seminal vesicles. 

Monocystis lloidi Seminal vesicles. 

Monocystis pheretimi . . Seminal vesicles. 

Stomatophora diadema. Seminal vesicles. 

Dirhynchocystisglobosa. Seminal vesicles. 


•Ozobranchus shipleyi Haemogregarina nicorise Body. 

Dendrostoma signifer .... Extreniocystis dendro- Ccelomic cavity. 



Methods for the examination and study of Protozoa are 
a.dequately dealt with in such works as Prowazek and Jollos 
(1921), Wenyon (1926), Hartmann (1928), Belaf (1928), 
Oatenby and Cowdry (1928), Knowles (1929) and Hegner 
and Andrews (1930). The principal methods followed in the 
study of the Sporozoa are given here for the benefit of those 
taking up the study of the group. 

Examination in the Living Condition. — It is always desirable 
to make observations on the hving organisms in the first 
instance. The specimens are mostly studied in a drop of the 
natural medium or body-fluid of the host -animal in which 
they are found. Any pressure of the cover -glass might cause 
deformities, and this should be guarded against by including 
in the preparation small bits of detritus or a hair which will 
support the cover -glass. If it is intended to continue the 
observations for some length of time, " cavity slides " may be 
used. The drop of fluid containing the organisms should be 
put on a circular cover-glass, the margin of which is smeared 
with melted paraffin or Czokor's wax, and the cover-glass 
inverted over the cavity. This will fix the cover-glass to the 
shde and prevent evaporation. The wax is made by heating 
together and mixing in a shallow tin provided with a lid 
■equal weights of bees-wax and Venetian turpentine. The 
wax becomes a soHd mass when cool, and can be appHed 
by placing the heated portion of a wire on the wax and then 
passing it round the cover -glass or the slide. To study the 
developmental stages and natural movements the slide should 
be kept at the body temperature of the host when this is 
a warm-blooded animal. 


Oocysts of CocciDiA escaping in the faeces in an incompletely 
developed condition may be observed to complete their 
development, especially when mixed with a 5 per cent, 
solution of bichromate of potash. Malarial parasites, when 
observed as described above, Avill continue to develop for 
several hours, and the production of merozoites can sometimes 
be observed. Occasionally the removal of the blood from 
the Vertebrate host may induce the development of stages 
which normally take place inside the body of an Invertebrate 
host : thus the production of gametes from the gametocytes, 
fertihzation, and the development of ookinetes may be followed 
in a drop of blood enclosed between a sHde and a cover -glass. 
During the observations the cover-glass may be removed and 
permanent preparations made showing particular stages of 

When the body-fluid containing the parasites is too small in 
amount to make a suitable preparation, physiological salt 
solution (0-7 per cent, solution of sodium chloride in distilled 
water for cold-blooded animals and 0-85 or 0-9 per cent, 
solution for warm-blooded animals) or Ringer's solution may 
be added. Ringer's solution consists of sodium chloride 
0'8 grm., potassium chloride 0-02 grm., calcium chloride 
0-02 grm., sodium bicarbonate 0-02 grm., distilled water 
100 c.c. 

Intra-vitam Staining. — Examination of the hving organisms 
is facihtated by intra-vitam staining, different parts of the 
organism or its contents being coloured without kilHng the 
animal or affecting its movements. For this purpose neutral 
red or methylene blue are used in very dilute solutions 
(1 in 10,000). A drop of the stain may be allowed to dry 
on the slide and the fluid containing the organisms be placed 
on the area and covered with a cover-glass. The stains are 
slowly absorbed by the parasites. Neutral red assumes 
a bright cherry-red colour in acid and a brown colour in 
alkaline media, and thus serves to indicate the reaction of the 
substance which it stains. 

Eosin is used as an indicator of the life or death of an 
organism. It will not stain hving cytoplasm, and by running 
a dilute solution under the cover-glass the Uving organisms 
can be distinguished from those that are dead. 

Cultivation. — Various methods have been adopted to provide 
favourable conditions of growth and multiphcation, so as to 
afford large numbers of a particular organism outside the body 
of the host for extensive study or for the purposes of a class. 
No artificial medium has so far been discovered in which 
Gregarinida or Coccidia will grow : but various methods 
have proved successful for the cultivation of blood-inhabiting 
parasites, as the malarial parasites and the Piroplasmids. 


These cultures are incubated at 37° to 40° C, and the forms 
which appear are those which normally occur in the Vertebrate 
host. Bass and Johns (1912) were the first to succeed in 
cultivating malarial parasites. Their methods, with shght 
modifications, were successfully followed by Ziemann (1913, 
1914), J. G. & D. Thomson, (1913), Row (1917), Suiton (1922), 
and others. Knuth and Richters (1913), Ziemann (1913, 
1914), and Thomson and Fantham (1914) were the first to 
apply these methods to the Piroplasmids. The technique of 
some of these methods is given below : — 

Bass and Johns's Method. — Withdraw 10 c.c. of malarial blood 
from a vein with a syringe, and transfer immediately to a de- 
fribinating flask containing 0-1 c.c. of 50 per cent, solution of 
dextrose. Defibrinate by stirring with a glass rod. Transfer 
the defibrinated dextrose blood to culture-tubes not less than 
1-25 cm. in diameter and 12-5 cm. in depth. The quantity 
of blood in each tube may vary in depth from 2-5 to 5 cm., 
which will give a column of serum 1-25 to 2-5 cm. deep above 
the cells, when they have settled to the bottom of the tube. 
Incubate the tubes in the vertical position at a temperature 
of 40° C. The parasites live and develop in the red blood- 
corpuscles just below the surface of the deposit. Carefully 
withdraw the cells from this layer by means of a fine pipette 
and examine at intervals. The young trophozoites will be seen 
to grow into schizonts and break up into merozoites. Some 
of these may enter other corpuscles and grow into schizonts, 
but as a rule those escaping from the corpuscles are devoured 
by the leucocytes. To remove the leucocytes centrifuge 
the defibrinated glucose blood at a speed sufficient to cause the 
leucocytes to occupy the upper layer of the deposit. Transfer 
the supernatant serum to flat-bottomed culture-tubes, fllhng 
to a depth of 1-25 to 2-5 cm. Pass a pipette into the middle 
of the deposit in the centrifuge -tube, draw off the red corpuscles 
with the parasites, and transfer them to the bottom of the 
culture -tubes. In this way, in the absence of the leucocytes, 
the parasites may complete two or three cycles ; but it has not 
been found possible to cultivate them for more than three 
generations. For making subcultures, take normal blood, 
treat it in the same manner, and inoculate it by means of 
a pipette with infected blood from the previous culture. 

Row (1917) and Sinton (1922) have devised modifications 
by which the growth of a single generation can be followed with 
a few drops of blood. 

Row's Method. — Draw the blood from the finger into a smaU 
tube and defibrinate it in the same. Transfer the small 
quantity of defibrinated blood by means of a pipette to a small, 
flat-bottomed tube containing serum to which the requisite 
quantity of dextrose has been added. Place the small tube 


in a larger tube (the ordinary bacteriological potato -tube) 
provided with a constriction on which the smaller tube may 
rest. In the portion of the outer tube below the constriction 
put some solution of pyrogallic acid and add 2 or 3 c.c. of 
•a 10 per cent, solution of sodium hydrate immediately before 
introducing the smaller tube. Cork the larger tube tightly 
with a rubber cork. The pyrogallic acid and sodium hydrate 
absorb the oxygen, and the culture thus takes place in an 
oxygen-free atmosphere. 

Sinton's Method. — A specially constructed tube about 20 cm. 
in length is employed (fig. 2). To prepare this apparatus, 
take a tube of 0-4 to 0-5 cm. bore. Draw out one end as in 
an ordinary pipette and, shpping a narrow metal tube over 
the thin drawn-out portion while it is still soft, press upwards 
to produce a dilated bulb with its lower surface flattened (B) 
a,nd with the thin drawn-out tube arising from it (A). Now 
Jieat the tube a short distance, about 1 cm., above the flattened 
surface and draw out till it forms a tube about 6 to 8 cm. 
long and 0-2 cm. wide (C). At the upper end of this narrow 
tube make a sHght constriction (D), and about 0-4 to 0-5 cm. 
■above it make another (E). Drop three glass beads (F) 
into the upper part of the tube (G), which is allowed to remain 
wider in diameter, and then draw out and bend as in a Wright's 
capsule (H). Keep the upper and lower drawn-out ends sealed 
and sterihze the whole tube. 

To make the culture, open the tube at both ends, and bend 
the upper capillary portion at right angles to the rest, so that 
the apparatus may lie on the table with the open upper end 
pointing upwards. Insert the upper end into an already 
prepared Wright's capsule containing ascitic or hydrocoele 
fluid, to which the requisite quantity of dextrose has been added, 
and allow the fluid to enter till the upper section is about a third 
or half full. Prick the carefully steriHzed finger of a malarial 
patient, and allow five to ten drops of blood to run into 
the fluid in the tube. Now gently heat the dilated part of the 
lower end of the tube and seal off the narrow part below it. 
The air in the dilated part will cool and the blood-mixture will 
be drawn further into the tube. Now seal ofl" the upper end 
also. Defibrinate the mixture by shaking the glass beads, 
and when this has been completed, swing round the tube 
rapidly so as to drive the defibrinated blood-mixture through 
the constriction into the lowermost part of the tube, so that 
it fills the dilated part and the narrow section above it. 
Heat the tube at the constriction above the column of fluid 
and seal ofl". The red corpusc]es will settle to the flat bottom. 
Incubate the tube in a vertical position at a temperature 
of 35° to 38° C. In order to examine, open the tube, withdraw 
by a pipette the cells from the lower end, and seal again. 



Maintenance of Parasitic Protozoa in Laboratory Animals. — 
In a laboratory strains of parasitic Protozoa may be main- 
tained in suitable animals which have been found by previous 
examination not to harbour natural parasites of their own. 
This method serves the same purpose as cultivation, and it is 
easier to maintain a strain in an animal host than in a culture. 
Blood parasites, such as the malarial parasites and the piro- 
plasmata, can be introduced by inoculating the blood from 
an infected to a clean host, and parasites that pass out in the 






Fig. 2. — Sinton's apparatus for cultivation of malarial parasites in 
a small quantity of blood. The smaU Wright's capsule 
shown on the right contains the ascitic or hydrocoele fluid. 
(After Sinton.) 

encysted condition, such as Coccidia, can be mixed with the 
food and allowed to be ingested. The practice has been 
particularly successful in certain special cases. 

The malarial parasites of birds can be maintained for a long 
time in canaries, sparrows, and other susceptible birds by 
inoculating blood from the one to the other. Some birds 
acquire heavy and fatal infections and others mild ones from 
which they recover. The piroplasmata of dogs are readily 


inoculable from one dog to another, but as fatal infections 
are often obtained, a number of animals will be required. 
Ticks fed on a dog will remain infective for long periods, and 
the parasites can thus be maintained in them. Strains of 
HsRmojproteus columhse can be maintained in pigeons by 
breeding Lynchia maura in the cages in which they are kept. 
A fresh pigeon is introduced into the cage from time to time 
to hand over the infection from fly to fly. Toxoplasma gondii 
is also inoculable to a variety of laboratory animals. 

Permanent Preparations of Fixed and Stained Protozoa. — 
Although it is important to study the hving organisms in the 
first instance, the finer details of structure can only be studied 
in properly fixed and stained smears or films. Some of the 
methods of general apphcation are described below. 

Staining under the Cover-slip. — For staining large organisms, 
such as the Gregarines, the following method is used : — 

Put wax feet at the corners of a square cover-shp, and invert 
it over a drop of the fluid containing the organisms. The wax 
feet should hold the cover-sHp firmly to the slide. With 
a pipette run a Httle fixative at one side of the cover-slip, and 
draw it through by holding a piece of filter-paper at the opposite 
side. When the fixative has had time to act, wash it out by 
substituting another fiuid (alcohol or water, as the case may be) 
and draw it through with filter-paper in the same manner. 
The stream should not be so violent as to wash away the 
organisms, but the substitution should be complete. Then 
run in the stain, allow it to act, and wash out and differentiate, 
if necessary controUing the process under the microscope. 
Dehydrate and then clear in clove-oil or xylol, and run in a very 
fluid Canada balsam . It is very important to see that the trans- 
fer from one fiuid to another is not too rapid, as otherwise 
there is great risk of shrinkage, and also to see that the dehydra- 
tion is complete. 

The following is an indication of the length of time generally 
required, but should be regarded as no more than an indication. 
Fix in Bouin's fluid, 5 min. ; wash in 70 per cent, alcohol, 
5 min. (several changes) ; stain in borax- carmine, 5 min. or 
more ; dehydrate with 70 per cent., 90 per cent., and absolute 
alcohol, 5 min. each, changing the absolute alcohol once or 
twice ; clear by running in a mixture of clove-oil and absolute 
alcohol and then pure clove-oil ; mount in Canada balsam 
by running the same under the cover-shp. 

Preparation of Wet Fixed Films. — Films made on shdes or 
cover-shps may be fixed with one of the fixing fluids, without 
being allowed to dry. After fixation the films are washed 
free of fixative, and stained and mounted Hke sections fixed 
on slides. The results shown by wet fixed films are far 
superior to those shown by dried films, described later. If the 


material to be examined is thick, it should be emulsified with 
physiological salt solution. In the case of blood, fluid from 
blood or serum culture media, tissues, faeces, in fact any 
hquid containing albuminous matter, the film will stick to 
the sUde or the cover-shp during the processes of fixation and 
staining. The usual fixatives and stains are described here- 

Staining the Organisms fixed on a Cover-slip. — ^Make a number 
of smears of the fluid containing the organisms on cover-sHps, 
and when the fluid, has partially dried, invert the cover-shps 
and let them float on the surface of the flxative contained in 
a dish ; or put a small drop of fluid containing the organism 
on a cover-shp and add with a pipette twice the quantity 
of hot fixative. When the organisms have been fixed on the 
cover-shps, pass successively through the washing fluid, 
alcohols, stain, clearing-fluid, etc., all these reagents being 
contained in shallow dishes. In all these subsequent stages 
put the cover-shps at the bottom of the fluid in the dish, 
with the face bearing the organisms upwards. Finally, 
remove the cover-shps and put them, face downwards, on 
shdes on each of which a drop of Canada balsam has been 
put. In this way quite a large number of smears or preparations 
can be fixed and stained in practically the same time as would 
be taken to make one preparation. 

Fixatives. — The foUowing are commonly employed for fixing 
the Protozoa : — 

(1) Concentrated solution of mercuric chloride, hot or 
cold : (2) Schaudinn's subHmate alcohol (2 parts saturated 
aqueous solution of mercuric chloride, 1 part absolute alcohol ; 
immediately before use, add acetic acid to the quantity to 
be used to the strength of 5 per cent.) : (3) Zenker's fiuid 
(mercuric chloride 5 grm., potassium bichromate 2-5 grm., 
sodium sulphate 1 grm. in 100 c.c. of distifled water, with 
2-5 to 5 per cent, of glacial acetic acid added before use) : 
(4) Bouin's fluid (saturated aqueous solution of picric acid 
75 parts, formol 25 parts, and acetic acid 5 parts) : (5) Bouin's 
alcohoHc fixative (picric acid 1 grm., 80 per cent, alcohol 
150 c.c, formol 60 c.c, acetic acid 15 c.c) : (6) formahn : 
and (7) vapour of 4 per cent, solution of osmic acid. Fixation 
is usuaUy complete in 15 to 30 minutes. 

Bouin's fixatives are the simplest to use, as the picric acid 
is more easfly washed out after fixation than the mercuric 
chloride contained in the others. After Bouin's fixative, 
wash the films in several changes of distilled water till all the 
picric acid has been removed. In the case of alcohohc Bouin, 
if an aqueous stain is to be used, commence washing in 
70 per cent, alcohol and, passing the film through graded 
alcohols, bring down to distilled water. After Schaudinn's 


fluid, wash in several changes of 70 per cent, alcohol to remove 
the mercuric chloride ; then wash in 70 per cent, alcohol to 
which a few drops of Wiegert's iodine solution have been 
added ; finally, wash in 70 per cent, alcohol to which a drop 
of 1 per cent, solution of sodium thiosulphate has been added, 
in order to remove all traces of iodine. Bring down through 
successive grades of alcohol mto distilled water if it is desired 
to use a watery stain ; a stay of a few minutes in each solution 
will sujB&ce . After fixation with Zenker's fluid, wash successively 
with distilled water, distilled water with a few drops of iodine 
solution, distilled water with a trace of 1 per cent, solution 
of sodium thiosulphate, and distilled water. 

Staining Methods. — The following stains are usually employed 
for staining Protozoa : — 

Borax-carmine. — Fix in Bouin's fluid for 10 to 20 minutes 
according to bulk and permeabihty. Wash out in 70 per cent, 
alcohol (several changes). Stain in borax-carmine till 
thoroughly penetrated ; 15 minutes are usually enough 
for small objects. Differentiate in acid alcohol (70 per cent, 
alcohol to which hydrochloric acid is added to the strength 
of 1 per cent.),controlhng under the microscope. Dehydrate 
in 90 per cent, to absolute alcohol. Put in a mixture of clove - 
oil and absolute alcohol, equal parts. After a few minutes 
transfer to pure clove-oil, and leave there till cleared. Mount 
in Canada balsam. 

Delafield's Hematoxylin. — Fix in Schaudinn's fluid. Wash 
in 30 per cent, alcohol, and bring down through 10 per cent, 
alcohol to distilled water. Add a few drops of Delafield's 
hsematoxylin solution to a watch-glass full of distiUed water. 
Leave in stain for a few minutes to an hour or more according 
to bulk. Bring up from distilled water through ascending 
grades of alcohol to 70 per cent, alcohol ; differentiate in 
70 per cent, acid alcohol. Dehydrate ; clear ; mount. 

Heidenhain's Iron Hsematoxylin. — Fix in Schaudinn's fluid 
for 10 to 30 minutes, according to the size and permeabihty 
of the object. Bring down to 30 per cent, and 10 per cent, 
alcohol to distilled water. Wash out the fixative thoroughly. 
Transfer to 4 per cent, solution of the violet crystals of iron 
alum (sulphate of iron and ammonium) in distilled water for 
\ hour to 12 hours according to the size of the organisms. 
Stain in Heidenhain's aqueous hsematoxylin solution (about 
0-5 per cent.) for 30 minutes to several hours. Wash in distilled 
water. Differentiate in 1 per cent, iron alum solution till 
the granules in the nuclei are distinct, the films being removed 
into distilled water and examined on a sHde with the micro- 
scope from time to time. When differentiation is complete, 
wash in several changes of distilled water and then for half 
an hour or more in running tap-water, Counterstain, if 



desired, with a 1 per cent, solution of eosin or orange G. 
Dehydrate by bringing through graded alcohols to absolute 
alcohol. Clear in xylol. Mount in Canada balsam. 

DobelVs Iron Hsematein. — Fix in Schaudinn's fluid. Bring 
down through 30 per cent, and 10 per cent, alcohol to distilled 
water. After washing, bring up through various grades of 
alcohol to 70 per cent., and from that transfer to 1 per cent, 
solution of iron alum in 70 per cent, alcohol for 10 minutes 
(the solution is made by dissolving 1 grm. iron alum in 23 c.c. 
warm distilled water and adding 77 c.c. of 90 per cent, alcohol). 
Rinse in 70 per cent, alcohol. Stain in 1 per cent, solution 
of hsematein in 70 per cent, alcohol for 10 minutes. Rinse 
in 70 per cent, alcohol. Differentiate films in original alum 
solution and sections in 70 per cent, acid alcohol. Wash in 
several changes of 70 per cent, alcohol. Dehydrate ; clear ; 
mount. The whole process may be carried out in 30 minutes. 
Light green in 90 per cent, alcohol may be used as a counter- 

Mayer's Hssmalum. — To prepare the stain dissolve 1 grm. 
of haematin in 50 c.c. of 90 per cent, alcohol, dissolve 50 grms. 
of alum in 1000 c.c. of water, and mix up the two solutions. 
Keep a crystal of thymol in the bottle to prevent growth 
of fungi, etc. 

When using the stain, add a few drops of the stain to a Petri 
dish of distilled water, and leave the films in for several hours 
or overnight. Wash the films in running tap -water till they 
are quite blue. Dehydrate by bringing up though various 
grades of alcohol to absolute alcohol. Clear in xylol and 
mount in Canada balsam. 

Mayer's Acid Hsemalum. — This strain is prepared from 
Mayer's hsemalum by adding acetic acid to a strength of 
2 per cent. It is used in the same manner, but has less 
tendency to overstain. 

If overstaining occurs with either of the above stains, 
place the films in acid alcohol. After they are decolorized, 
wash the films well in running water till the acid is completely 
neutrahzed and they are blue. 

Mallory's Eosin and Methylene Blue. — This is recommended 
for sections of tissues containing parasites. Fix in Zenker's 
fluid. Wash out the fixative in rumiing water for several 
hours. Stain in warm 5 per cent, aqueous solution of eosin 
for 20 minutes or longer, wash in water, and stain in Unna's 
alkahne methylene-blue solution diluted with 4 or 5 parts of 
water for 10 to 15 minutes. Wash in water, diffierentiate in 
95 per cent, alcohol, controlling under a microscope, until 
sections are pinkish but nuclei deep blue. Dehydrate quickly 
and mount. 

Mallory's Triple Stain. — ^This is also recommended for 


sections. Fix in Zenker's fluid. Thoroughly wash out the 
fixative for several hours in gently running water. Stain 
sections in 0-5 per cent, aqueous solution of acid fuchsin for 

2 to 4 minutes, and transfer to the second solution (consisting 
of aniline blue soluble in Avater (Griibler) 0-5 grm., orange G 
(Griibler) 2 grm., 1 per cent, aqueous solution of phosphomo- 
lybdic acid 100 c.c.) for 10 to 20 minutes or longer. Wash 
and differentiate the sections in tap-water, dehydrate rapidly, 
clear, and mount. Lund (1933) recommends Zenker's fluid 
with 2-5 per cent, acetic acid, staining the sections for 2 minutes 
in Mallory No. 1 and 1 minute in Mallory No. II, then dipping 
rapidly into 95 per cent, and absolute alcohol, blotting quickly 
between each change, and then clearing in xylol for about 

3 minutes. 

Preparation of Dried Films on Slides. — Films of blood, in- 
testinal contents or other fluids containing parasites are usually 
made for diagnostic purposes. Films of sohd organs, such 
as the Kver or the spleen, are made by smearing a piece of the 
organ hghtly across the shde or dabbing the slide with the 
freshly-cut surface. The film is dried as rapidly as possible 
by waving it in the air. Distortion and disintegration of the 
parasites may take place in the process of making the films 
and subsequent drying ; to avoid this expose the film to 
osmic acid vapour for ten to fifteen seconds before allowing 
it to dry. 

Thin Blood-films. — To make blood-films take perfectly clean 
glass-shdes, free from any trace of grease, and lay them flat 
on a clean piece of paper. Sterilize a needle in the flame, 
and prick the dorsum of a finger just below the root of the nail. 
Pick up one of the clean shdes, invert it, and towards one end 
just bring it in contact with the oozing blood. Too large 
a drop must not be taken. Re-invert the shde on the table 
and, taking another slide, with a smooth even edge as a spreader 
hold in at an angle of 45°, touching the drop so that a thin 
film of blood runs between the edge of the spreader and the 
shde. When the blood has spread along the edge, push the 
spreader fairly rapidly toward the other end. The blood must 
follow the spreader and not be pushed before it. A thin 
film will result covering about one-half of the shde. Allow 
the film to dry in air, covering it, if necessary, with a Petri 
dish to protect it from flies, dust, etc. In monsoon weather 
haemolysis may take place before the film has time to dry, 
and at such times the film should be held, as soon as made, 
over a spirit-lamp flame to dry. Films are usually stained 
by one of the modifications of Romanowsky's stain. 

Staining the Dried Blood-films. — The films, as prepared 
above, may be stained with the original Romanowsky's stain, 
or with one of the many modifications, such as Leishman's 


stain or Giemsa's stain. The action of all such stains depends 
on certain loose chemical combinations. Medicinal (not 
pure) methylene-blue contains a number of oxidation products, 
the most important of which is methylene-azur. When watery 
or alcohohc solutions of methylene-blue and of eosin are mixed 
together a series of loosely combined chemical bodies are formed. 
These different compounds possess different affinities for 
different cell-structures, and thus differential staining results. 
The red blood- corpuscles stain a transparent pink or orange 
colour ; the nuclei of leucocystes, shades of violet ; eosinophile 
granules in the coarsely granular leucocytes, red ; the cyto- 
plasm of malaria and other blood-inhabiting protozoal parasites 
a bright " Cambridge " blue, and their chromatin a bright 
ruby -red. 

The Original Eomanowsky's Stain. — In large laboratories the 
original Romanowsky's stain is still employed, as it is more 
economical than Leishman or Giemsa. Two stock solutions are 
needed. For preparing Solution A take medicinal (not pure) 
methylene-blue 1 grm., pure sodium carbonate 0-5 grm., 
distilled water 100 c.c, dissolve and place the stain either in 
the 37° C. incubator or in full sun-Hght for 2 or 3 days. The 
solution should acquire a deep purple colour before it is ready 
for use. For Solution B, dissolve eosin, extra B.A., water 
soluble 0-1 grm. in 100 c.c. of distilled water. 

Fix the film for 5 minutes in methyl alcohol, or for 10 minutes 
in ordinary alcohol. Wash with distilled water and transfer 
to a clear Petri dish. Dilute 1 part of solution A with 19 parts 
of distilled water. Similarly dilute 1 part of solution B with 
19 parts of distilled water. Mix equal parts of the diluted 
solutions A and B, and pour the mixed stain, immediately 
after mixing, into the Petri dish to cover the shde. Stain for 
half an hour or longer. On tilting the Petri dish a red stain 
should be seen at the edge, which will indicate that the 
staining is proceeding properly. Wash off the stain with 
distilled water and differentiate in a bath of distilled water. 
When the film commences to turn pink, remove from the 
bath and allow it to dry by slanting it against a vertical 

Leishman' s Stain. — To prepare the stain thoroughly clean 
a glass (not porcelain) pestle and mortar and rinse out the 
mortar with a httle pure methyl alcohol. Weigh out 0-15 grm. 
of Leishman's stain powder (Merck's or Griibler's). Measure 
into a perfectly clean glass cylinder 100 c.c. of methyl alcohol, 
purissimum, acetone-free (Merck's). Add a httle of the alcohol 
to the stain in the mortar and grind. Add more alcohol and 
grind. Drain off the dissolved stain into a stoppered bottle. 
Continue to add the alcohol in portions and grind until every 
particle of the stain has gone into solution, and use the fuU 


100 c.c. of methyl alcohol. Next ripen the stain by placing 
the bottle of stain overnight (but not longer) in the 37° C. 

In order to stain, lay the blood-film, film side upwards, on 
a staining rack. Drop the stain on to the sHde until the 
whole surface is covered with the stain. The methyl alcohol 
in the stain fixes the film. This takes half a minute only ; 
after that interval drop on to the shde double the number of 
drops of pure distilled water. By tilting the end of the shde 
allow the water and stain to mix thoroughly, and allow the stain 
to act for 5 minutes or more. Wash the stain ofi" by immersing 
the slide in distilled water. Put the shde to soak in a clean 
Petri dish containing fresh distilled water and rock gently. 
The film turns at first greenish-blue, then pink. This takes 
about half a minute or less. When the film is just turning 
pink remove the slide and lean it against a vertical surface 
to dry. 

Giemsa's Stain. — To prepare the stain grind in a glass mortar 
with a glass pestle 3 grm. of azur-II-eosin and 0-8 grm, of 
azur-II (Griibler's or Merck's) into thorough solution in 250 c.c. 
of the purest anhydrous glycerine. Add 250 c.c. of the purest 
acetone-free methyl alcohol and mix thoroughly. Allow to 
stand overnight and next day filter the stain through filter 
paper. Prepared Giemsa's stain is stocked by many firms, 
and is also supphed by the Central Research Institute, Kasauli. 

Fix the film by covering it for 3 to 5 minutes Avith pure 
methyl alcohol or by dipping it for 10 minutes into absolute 
alcohol, and wash thoroughly in distilled water. Dilute 
1 part of Giemsa's stain with 14 parts of distilled water 
(10 or 15 drops in as many c.c. of distilled water), and pour 
it over the slide placed in a Petri dish. Stain for half an hour 
or longer. Remove the slide, flush with distilled water, and 
put it in a bath of fresh distilled water until it commences to 
turn pink. Remove the slide and let it dry by leaning it 
against a vertical surface. 

The Panoptic Method of Staining. — Lay the shde on a stain- 
ing rack and cover it for half a minute with undiluted Leish- 
man's stain ; dilute the stain with double the number of drops 
of distilled water, and allow to stain for 5 or 10 minutes. 
Wash the film with distilled water. Then lay the shde in 
a Petri dish and flood with diluted Giemsa's stain, one drop to 
each c.c. of water. Stain for 1 to 24 hours, covering the 
Petri dish to prevent evaporation. Wash the fihn with distilled 
water and transfer it to a bath of 1 in 1000 acetic acid. When 
the film begins to turn pink, remove, wash rapidly with dis- 
tilled water, and slant the slide against a vertical surface to dry. 

A more rapid alternative method is as follows : — Cover the 
film with undiluted Leishman's stain for half a minute ; 


dilute the stain with double the volume of diluted Giemsa's 
stain, one drop to each c.c. of water, and mix thoroughly. 
Differentiate in distilled water, as in the ease of Leishman's 

Thick Blood-films. — The thick film method is specially 
applicable to those cases in which the patient has already taken 
a small dose of quinine and the malarial parasites are apt to 
be missed in thin films. Knowles (1928) recommends taking 
three thin films and one thick one from every suspected case 
of malaria. The thin films are stained and examined whilst 
the thick film is drying, and if no parasites are detected in the 
thin films the thick film may be proceeded with. Knowles 
and Das-Gupta (1924 a) advocate the following method for 
preparing thick films : — Take a perfectly clean shde, prick the 
patient's finger, and bring the slide into contact with the 
issuing blood so that four drops are placed at the corners of 
a small square about half an inch across. The drops should 
not be too large nor too small. With a round needle pool 
the four drops so as to spread the film into an even thick film 
covering the half -inch square. Puddling should be avoided 
and the film must not be made too thick. The film should be 
kept covered, and will take about two hours to dry. Lay the 
shde on a staining rack and dehsemoglobinize the film by 
gently flooding the shde with a mixture of 2-5 per cent, 
solution in distilled water of glacial acetic acid, 4 parts, and 
2 per cent, solution in distilled water of crystalline tartaric 
acid, 1 part. The dehsemoglobinized film should have a grey- 
white colour. As soon as the process is complete drain off 
the fluid by gently tilting the shde. Next flood the shde with 
methyl alcohol and allow it to remain on for 3 to 4 minutes. 
Drain off the methyl alcohol and wash the film thoroughly in 
distilled water so as to remove every trace of acid. Stain with 
diluted Giemsa's stain, one drop to each c.c. of water, for 
15 minutes or longer. Differentiate in the usual way with 
distilled water. Do not blot the film, but let it dry by 
slanting it against a vertical surface. 




Class SPOROZOA Leuckart, 1879. 

Protozoa which are exclusively parasitic in their mode of 
life, and Uve in the cells or body fluids of Vertebrate or Inverte- 
brate hosts. They produce resistant spores at some stage 
or another of their hfe- cycle. 

The general organization of the class and the basis of classi- 
fication into subclasses and orders has already been discussed. 
Following Reichenow, the class Sporozoa is divided into 
four subclasses, as follows : — 

1 (2). Trophic and reproductive phases typi- 
cally distinct ; trophozoite becoming 

fully developed before reproduction [Schaudinn, p. 50. 

begins. Sporozoites are Gregarinulse . Telosporidia 

2(1). Trophic and reproductive phases usually 
overlap ; the still-growing or even quite 
yoiuig trophozoite may begin to form 
spores. Sporozoites are Amcebulse ... 3. 

3 (4). Spores with one or more thread-cap- [p. 328. 

sules Cnidosporidia Doflein, 

4 (3). Spores without thread-capsules 5. 

5 (6). Cysts forming long rod-like masses 

(Miescher's tubes). Spores crescentic, 
with one end rounded and the other 
pointed. Parasites of striped muscle [p. 361. 

of Vertebrates Sarcosporidia Balbiani, 

6(5). Spores large, containing a single voliimi- 

nous nucleus. Simple type of develop- [& Mesnil, p. 369. 

ment Haplosporidia CauUery 

I. Subclass TELOSPORIDIA Schaudinn, 


Telosporidia are Sporozoa in which the trophic and 
reproductive phases are typically distinct, the animal becoming 
full grown before either asexual reproduction {schizogony) or 
sexual reproduction {gamogony or sporogony) begins. 

The great majority are intracellular parasites, at least 
during part of their life-history. The parasites are uninucleate 
in the early period of growth. During growth (as in Coccidia 
and H^mosporidia). or after growth is completed (as in 
Gregarinida), the nucleus divides by repeated mitotic 
divisions, and the organism breaks up into as many daughter 


individuals as the number of nuclei. This process of multiple 
j&ssion may be an asexual process {schizogony), and the resulting 
daughter individuals (merozoites) will grow into adult tropho- 
zoites, or the daughter individuals may all be gametes. In 
the former case also, after schizogony has been repeated 
a number of times, some of the merozoites develop into 
gametocytes, which give rise to gametes. The gametes 
conjugate, the process being known as syngamy, and form 
zygotes. The zygote then becomes enclosed in a resistant 
cyst, known as the oocyst. Inside the oocyst the zygote 
may either divide directly into a number of motile vermiform 
bodies called sporozoites, or the zygote may &st divide into 
separate bodies called sporoblasts, which become encysted in 
secondary cysts known as sporocysts ; within each sporocyst 
the sporoblast gives rise to a number of sporozoites and 
a residual body. The mature oocyst of this latter type thus 
contains a number of sporocysts, each containing many 

The infection of a new host is brought about by the con- 
taminative or inoculative method, and the sporozoites seek 
their way to the particular type of cell which they usually 
parasitize, viz., an epithehal cell (Gregakinida, Coccidia) 
or a blood-corpuscle (H^mosporidia) . The adult forms of 
CrREGARiNiDA are invariably extracellular or lumen dwelHng, 
young growing stages alone being intracellular. Adult forms 
of Coccidia and H^mosporidia are persistently intracellular, 
young, adult, and reproductive phases all occurring inside 
ahost-cell. In the majority of Gregarinida (Eugregarinaria), 
the sporozoite grows directly into a gametocyte which produces 
gametes. In the Schizogregarinida schizogony also takes 
place. In the Coccidia and H^mosporidia regular alterna- 
tion of the asexual method or schizogony, and the sexual method 
or sporogony, is the rule, and this alternation of generations 
frequently takes place in two distinct hosts. In Gregarinida 
and the H^mosporidia the sporozoites are formed directly 
by divisions of the zygote ; in Coccidia the zygote encysts 
and divides into sporoblasts which become sporocysts, and the 
oocyst thus contains a number of sporocysts, inside which the 
sporozoites are developed. When sporocysts are present, 
the oocyst is termed sporocystid, and, according to the 
number present, is described as disporocystid, tetrasporocystid, 
or poly sporocystid. The sporocysts are described as monozoic, 
dizoic, tetrazoic, octozoic, etc., according to the number of 
sporozoites present in each. 

The subclass is divided into three orders, as follows : — 

Mature trophozoite extracellular, large ; 

zygote non-motile ; sporozoites within [em. Doflein, p. 52. 

a spore Gregarinida A. Schneidei*, 




Mature trophozoite intracellular and small. 
Zygote non-motile ; sporozoites within 

a spore Coceidia Leuckart, p. 113. 

Zygote motile ; sporozoites without [sky, em. Doflein, p. 209. 
envelope Hsemosporidia Danilew- 

I. Order GREGARINIDA A.Schneider, 
emend. Doflein, 1901. 

The Gkbgaeinida are chiefly ccelozoic or lumen-dwelHng 
parasites of Invertebrates, especially Arthropods and Annehds, 
usually inhabiting the digestive tract, less frequently the 
ccBlome or the vascular system. They are typically intra- 
cellular only in the early part of their growth, i. e., in the 

Fig. 3. — Life-cycle of a typical Eugregarine. A, syzygy ; B, union 
of two sporonts ; C, encystment of the two sporonts and 
nuclear multiplication in each ; D, arrangement of the 
nuclei on the surface (pearl stage) ; E, gamete formation ; 
F, gametes set free and their union in pairs ; G, cyst con- 
taining the spores ; H, liberation of the spores through the 
sporoducts. (From Reichenow, after Rahler, Schnitzler, and 

trophozoite stage. Later they leave the epithelial cell and 
develop into more or less elongate motile adults, usually 
referred to as Gregarines. The vast majority of the Gregari- 
NiDA do not show asexual reproduction or schizogony, and 
multiphcation takes place solely by sporogony following 
upon gametogony. The adult Gregarines or sporonts are 
gametocytes, but do not show a differentiation into male or 
female gametocytes, which is a characteristic feature of the 
CocciDiA and the H^mosporidia. The gametocytes associate 


in pairs, and both produce an equal number of gametes, which 
are usually equal in size though not always in character, 
since those arising from one gametocyte may behave as male 
gametes and those from the other as female gametes. Con- 
jugation thus takes place between similar or dissimilar gametes 
{isogamy or anisogamy) . In a small group, however, schizogony 
takes place as well as sexual reproduction, and the order is 
divided into two suborders on this basis. Thus : 

No schizogony Suborder Eugregarinaria Doflein, p. 53. 

Schizogony takes place . . . Suborder Schizogregarinaria Leger, p. 111. 

I. Suborder EUGREGARINARIA Doflein, 1901. 

This suborder includes the majority of the so-called Gre- 
garines, which are usually parasites of Arthropods or Annelids. 
The sporocyst, after gaining entrance to a suitable host, 
germinates and sets free the sporozoites, which enter the 
epithehal cells of the digestive tract. There they grow large 
and protrude from the host-cells, remaining attached to them 
by an organ of attachment, called the epimerite, of varied form. 
These trophozoites sooner or later become detached from the 
ceUs of the host and move about in the lumen of the gut, 
where they are usually encountered as large and vermiform 
bodies exhibiting a ghding movement. In one group, the 
Sbptata, the main body is divided into two parts, which are 
distinctly marked off from each other by an ectoplasmic 
septum. The smaller anterior part is known as the protomerite, 
and the larger posterior part the deutomerite. The latter 
contains a single nucleus. In the other group, the Haplocyta, 
the body is not divided by a septum, and consists of a single 
chamber. In this latter group the sporozoites penetrate the waU 
of the gut and enter the body- cavity, forming cysts on the 
coelomic side of the intestinal waU, or develop as free forms 
inside the seminal vesicles or other parts of the body-cavity. 

The trophozoites increase in size and are then known as 
gametocytes ; these then encyst in pairs. Within the cyst- 
membrane each gametocyte gives rise to a large number of 
gametes, which may be isogamous or anisogamous. Each of 
the gametes formed from one gametocyte unites with one 
formed from the other, and a large number of zygotes are 
the result. Each zygote becomes surrounded by a resistant 
membrane (oocyst or sporocyst), and its contents produce 
usually eight sporozoites. 

The suborder is divided as follows : — 

[kester, p. 54. 

Trophozoite single-chambered Legion Haplocyta Lan- 

Trophozoite divided by an ectoplasmic [p. 89. 

septum Legion Septata Lankester, 


1. Legion HAPLOCYTA Lankester, 1885 

(=AcEPHALrN"A Koelliker or Monocystidea Stein+Lecudi- 
nidee and other primarily Dicystid forms). 

The Efgrbgabinaeia are usually divided into Acephatjna 
and Cephalina on the basis of the absence or presence of 
an epimerite. Hesse (1909), Bhatia (1924), and Cognetti 
(1921, 1925, and 1926) have shown that a fairly large number 
of Monocystids also possess an organ of attachment or epimerite. 
An epimerite is also present in the family Lecudinidse (=Dolio- 
cystidse), but the body is not divided into a protomerite and 
a deutomerite. Brasil (1908 and 1909) discussed the relation- 
ship of DoUocystis with Lankesteria and other admittedly 
Monocystid forms. Minchin (1903) considered it piu-ely 
a matter of definition whether Doliocystidee should be considered 
as Cephalina Avithout a septum, or as Monocystidea with 
an epimerite. I have elsewhere (1924, p. 508) discussed the 
grounds for classifying the Eugregarinakia into Haplocyta 
and Septata, and for placing the DoHocystidse under the 

The legion Haplocyta may be divided into two tribes, 
as follows : — 

1. Producing sporocysts with similar poles. 

Generally ccelomic parasites of Oligo- [p. 54. 

chsetes Homopolaridea Bhatia, 

2. Producing sporocysts with dissunilar poles. 

Intestinal or ccelomic parasites, usually of [Bhatia, p. 82. 

marine animals Heteropolaridea 

1. Tribe HOMOPOLARIDEA Bhatia, 1930 

(^Family Homopolarid^ Dogiel). 

Sporocysts with similar poles. Generally ccelomic parasites 
of terrestrial Oligochsetes. The tribe comprises eight famiHes 
{vide Bhatia, 1930), of which representatives of six are at 
present known from India. 

Identification Table of Families. 

1 (12). Sporocysts biconical, with similar, non- 

appendiculate poles, octozoic 2. 

2 (11). Sporocysts not provided with spines at 

either extremity 3. 

3 ( 10). Trophozoite simple 4. 

4 (9). Trophozoite solitary 5. [Bhatia, p. 55. 

5 (6). Trophozoite without any differentiation [Stein, emend. 

at the anterior end Monocystidae 

6 (5). Trophozoite with a differentiation at the 

anterior end 7. 

7 (8). Trophozoite with a conical or cylindro- [Bhatia, p. 65. 

conical trunk at the anterior end Bhynchocystidse 


8 (7). Trophozoite with a sucker-like differentia- [Bhatia, p. 68. 

tion at the anterior end Stomatophoridse 

9 (4). Adult trophozoites always associated in [Bhatia, p. 72. 

pairs Zygocystidae 

10 (3). Trophozoite branching at the anterior [Bhatia, p. 73. 

end Aikinetocystidae 

11 (2). Sporocysts provided with spines at either [Bhatia. 

extremity Syncystidse * 

12 (1). Sporocysts not bieonical. Trophozoites 

with or without an epimerite 13. 

13 (14). Sporocysts oval or spherical. Gametes [Bhatia, p. 76. 

net differentiated Diplocystidse 

14 (13). Sporocysts spherical. Gametes differenti- [Poche. 

ated Schaudinellidae *, 

1. Family MONOCYSTIDiE Stein, emend. 
Bhatia, 1930. 

Trophozoites oval, spherical or elongated, without any 
differentiation at the anterior end, sohtary. Sporocysts 
bieonical, with similar, non-appendiculate poles, octozoic. 
The family includes four genera {vide Bhatia, 1930), of which 
three are known from India. 

Key to Indian Genera. 

1 (4). Trophozoite ovoid or spherical 2. [p. 55. 

2 (3). Trophozoite ovoid Monocystis Stein, 

[p. 59. 

3 (2). Trophozoite spherical Apolocystis Cognetti, 

4(1). Trophozoite elongated, cylindrical, lil^e [p. 60. 

a Nematode worm Nematocystis Hesse, 

Genus MONOCYSTIS Stein, 1848, emend. Hesse, 1909, 
and Cognetti, 1923. 

Monocystis, Stein, 1848, pp. 183-223 ; Lankester, 1863, p. 93 
1866, pp. 23-8 ; 1872, pp. 342-51 ; Blitschli, 1880-2, p. 576 
Ruschhaupt, 1885, pp. 1-38 ; Bosanquet, 1894, pp. 421-33 
Labbe, 1899, p. 38 ; Cuenot, 1901, pp. 581-654 ; Cecconi, 1902 a 
pp. 132-5 ; 1902 b, pp. 122-40 ; Minchin, 1903, pp. 154-66 
Brasil, 1905 a, pp. 17-38 ; 1905 b, pp. 69-99 ; Hesse, 1909, p. 44 
Hoffmann, 1909, pp. 139-66 ; Mulsow, 1911, pp. 20-55 ; Minchin, 
1912, pp. 23, 174, 328, 331, 332, 333, 336, 339 ; Cognetti, 1923, 
pp. 250-3 ; Berlin, 1924, p. 14 ; Cognetti, 1925, p. 229 ; Wenyon, 
1926, p. 1147 ; Calkins & Bowling, 1926, pp. 385-99 ; Knowles, 
1928, p. 499; Reichenow, 1929, p. 886; Bhatia, 1929, p. 122; 
1930, p. 158 ; Kudo, 1931, pp. 57, 74, 293 ; Calkins, 1933, pp. 99, 
100, 309, 559. 

Trophozoites ovoid, without marked differentiation anteriorly^ 
often provided at the anterior pole with a mucron, solitary. 
Sporocysts as defined for the family. 

* placed after the name of a family indicates that no representative 
of the family has as yet been recorded from India, Burma or Ceylon. 



Key to Indian Species. 

1 (3). Nucleus spherical, with a large central 

karyosome 2. 

2 . Trophozoites fusiform or variable in form, 

often with a clear cylindrical process at 

one end, 100-150ju by 30-40 /x. Gameto- [p. 56. 

cysts spherical, 80-100 /x in diameter . . M. beddardi Ghosh, 

3 (1). Nucleus spherical or oval, with an irregu- 

lar or eccentric karyosome 4. 

4 (6). Nucleus with a large irregular karyo- 

some, consisting of several masses ... 5. 
5. Trophozoites oval, or elongated and 
club-shaped, 40-80 /x by 12-30 ^. 

Gametocysts irregularly hemispherical, [p. 57. 

70-80 fi in diaraeter M. bengalensis Ghosh, 

6 (4). Nucleus spherical or oval, with an 

eccentric karyosome 7. 

7 (8). Trophozoites rounded or oval when 

yotmg, fusiform when mature, lOOyn 

in length. Gametocysts spherical or [p. 58. 

oval, 84 ;n in diameter M. lloidi Ghosh, 

8(7). Trophozoites variable in form, attaining 

a size up to 220 /a by 50 ;u,. Gameto- [p. 58. 

cysts spherical, 80 /x in diameter M. pheretimi Bh. & Ch., 

1. Monocystis beddardi Ghosh. (Fig. 4.) 

fMonocystis beddardi, Ghosh, 1923, pp. 425-7, figs. 9-13. 
Monocystis beddardi, Bhatia, 1929, p. 123. 

Trophozoites elongately fusiform when young, fusiform 
or variable in form when mature. Body often with a clear 

Fig. 4. — Monocystis beddardi Ghosh. (After Ghosh.) 

cylindrical process at one end. Surface smooth. Ectoplasm 
very thin. Endoplasm highly granular, with paramylon 
grains. Change of shape involves the formation of a bulbous 
swelhng, which occupies a large portion of the body, with 

t prefixed to a reference indicates that the record of the species 
from India, Burma or Ceylon is based on it. 


rapid in-pouring of the paramylon grains into it, and the 
swelhng gradually disappears, to be formed again ; the 
parasite now assumes a rounded shape, with or without 
a narrow process at one or both ends. Nucleus spherical, 
with a large central karyosome. Gametocyst spherical, with 
hemispherical or irregularly oval gametocytes. Sporocysts 

Dimensions. — Trophozoite 100-150 /i in length and 30-40 /x 
in width ; gametocysts 80-100 ju, in diameter ; sporocysts 
12-5 /x in length. 

Remarks. — Dr. K. N. Bahl sent me some preparations of the 
contents of the seminal vesicles of EutypTiseus sp. In these 
are a large number of trophozoites and other stages of a Mono- 
cystid referable to M. beddardi Ghosh, There are, in addition, 
numerous specimens referable to DirJiynchocystis globosa 
(Bhatia & Chatterjee). Ghosh in his description of Jf . beddardi 
describes the parasite as assuming a rounded shape, with or 
without a narrow process at one or both ends. It is possible 
that he found specimens of D. globosa occurring along with 
M. beddardi and considered them merely as contracted stages 
of the latter. But the processes in Dirhynchocystis are clearly 
marked off from the body, and not as represented by Ghosh 
in his figs. 12 and 13. 

Habitat. — Seminal vesicles of Eutyphoeus nicholsoni 
(Beddard) : Bengal, Calcutta ; Eutyphoeus sp. : United 
Provinces, Lucknow. 

2. Monocystis bengalensis Ghosh. (Fig. 5.) 

■\Monocystis bengalensis, Ghosh, 1923, pp. 423-5, figs. 1-7. 
Monocystis bengalensis, Bhatia, 1929, p. 123. 

Trophozoites oval when young, later elongated and club- 
shaped. Anterior end wide and rounded. Posterior end 


Fig. 5. — Monocystis bengalensis Ghosh. A, trophozoite ; B, cyst 
with two sporonts. (After Ghosh.) 

narrow and rounded. Surface smooth. Ectoplasm very thin. 
Myonemes barely visible in stained specimens. Endoplasm 
highly granular, with large irregular-shaped paramylon 
grains. Nucleus rounded, with a large irregular karyosome, 



consisting of several compact masses. Movement slow and 
peristaltic. Gametocytes rounded or oval. Gametocysts 
irregularly hemispherical, apposed to each other by their 
flat surfaces. Sporocysts and sporozoites typical. 

Dimensions. — Trophozoite 40-80 /x in length, 12-30 /x in 
width ; gametocysts 70-80 ju. in diameter. 

Habitat. — Seminal vesicles of Pheretima posthuma (L. 
Vaill.) : Bengal, Calcutta. 

3. Monocystis lloidi Ghosh. 

■\Monocystis lloidi, Ghosh, 1923, p. 425. 
Monocystis lloidi, Bhatia, 1929, p. 123. 

Trophozoites rounded or oval when young ; somewhat fusi- 
form, about one-third longer than broad, when mature. 
Anterior end more rounded than the posterior. Surface 
smooth. Ectoplasm distinct and comparatively thick. Endo- 
plasm finely granular, with scattered, coarse granules. 
No distinct paramylon grains. Nucleus with a small eccentric 
karyosome. Movement brisk and peristaltic. Gametocysts 
spherical or oval. Sporocysts and sporozoites typical. 

Dimensions. — Trophozoite 100 ju. in length ; gametocysts 84 /x 
in diameter. 

Habitat. — Seminal vesicles of Pheretima posthuma (L. 
Vaill.) : Bengt^, Calcutta. Rare. 

4. Monocystis pheretimi Bhatia & Chatterjee. (Fig. 6.) 

■\Mo7wcystis pheretimi, Bhatia & Chatterjee, 1925, pp. 199-200^ 
figs. 20, 21, 28, 29. 
Monocystis pheretimi Bhatia, 1929, p. 123. 

Trophozoites variable in form, spherical, ovoid, eUipsoidal 

Fig. 6. — Monocystis pheretimi Bh. & Oh. (After Bhatia and Chatterjee.) 


or dumbbell-shaped, showing active movements. Para- 
glycogen grains few. Epicyte thin, without striations. 
Sarcocyte well developed. Myocyte fibrils dehcate. Endo- 
plasm excavated by vacuoles. Nucleus generally spherical, 
sometimes ovoidal, with a single spherical karyosome, placed 
eccentrically. Gametocysts spherical. Sporocysts and sporo- 
zoites typical. 

Dimensions. — Trophozoites attain a size up to 220 jx by 
50 fjL ; gametocysts 80 /i in diameter. 

Habitat. — Seminal vesicles of Pheretima posthuma (L. 
Vaill.) : Punjab, Lahore ; Bombay, Bombay. 

Genus APOLOCYSTIS Cognetti, 1923. 

Apolocystis, Cognetti, 1923, p. 253 ; 1925, p. 231 ; Reichenow, 
1929, p. 887 ; Bhatia, 1929, p. 124 ; 1930, p. 158 ; Troisi, 1933, 
pp. 336-42 ; Calkins, 1933, p. 559. 

Trophozoites spherical, without any marked principal axis, 
and without polar differentiation, solitary. Sporocysts as 
in the family. , 

5. Apolocystis matthaii (Bhatia & Setna). (Eig. 7.) 

^Monocystis matthaii, Bhatia & Setna, 1926, pp. 362-4, figs. 1-4, 25-27. 
Apolocystis matthaii, Bhatia, 1929, p. 124. 

Trophozoites most commonly spherical, sometimes ovoid 
or kidney-shaped, not covered by hairs. Epicyte thin and 
not showing any meridional striations. Sarcocyte hyaline 
and httle developed. Myocyte fibres not distinguishable. 
Endoplasm very dark, being finely granular, the alveoh 

Fig. 1 .—Apolocystis matthaii (Bh. & Set.). (After Bhat a and Setna.) 

compactly arranged and with large grains of reserve material. 
Nucleus spherical, with single spherical karyosome, placed 
eccentrically. Gametocysts elHpsoidal. 

Dimensions. — ^Trophozoites attain a size up to 238 /x ; 
gametocysts up to 468 /x in their longer diameter. 


Remarks. — Instances of solitary encystment and, more 
rarely, cysts containing three individuals were encountered. 
I have emended the spelling of the specific name to make it 
conform correctly to that of the person to whom the species 
is dedicated. 

Habitat. — Seminal vesicles of Megascolex trilobatus (Steph.) : 
Bombay, Bombay. 

Genus NEMATOCYSTIS Hesse, 1909. 

Nematocystis, Hesse, 1909, p. 45 ; Cognetti, 1918, p. 149 ; 1921 a 
pp. 153-4 ; 1921 b, pp. 552-4 ; Berlin, 1924, pp. 67-8, 74, 83-7 
Cognetti, 1925, p. 231 ; Bhatia & Chatterjee, 1925, pp. 193-7 
Bhatia & Setna, 1926, pp. 364-7 ; Reichenow, 1929, p. 887 
Bhatia, 1929, pp. 124-5 ; 1930, p. 158 ; Calkins, 1933, p. 559. 

Trophozoites elongated, cyhndrical, shaped like a Nema- 
tode worm, attaining even up to 5 mm. in length. Solitary. 
Sporocysts as in the family. 

Key to Indian Species. 

1 (6). Trophozoite tapering at one or both ends. 2. 

2 (3). Trophozoite tapering at either end. 

Nucleus long, fusiform, with two [p. 60, 

karyosomes iV. hessei Bh. & Ch., 

3 (2). Trophozoite with anterior end rounded, 

posterior end pointed 4. 

(5). Nucleus long, fusiform, with single karyo- [p. 62. 

some N. lumbricoides Hesse, 

5 (4). Nucleus elongated oval, with several [Bh. & Ch., p. 62. 

karyosomes N. plurikaryosomata 

6 (1). Trophozoite usually rounded at both 

ends 7. 

7 (8). Anterior pole without epimeritic denti- 

culations, posterior pole not covered 
by fine hairs. Nucleus usually in the 
middle of the body, with single karyo- [Set., p. 63. 

some N. stephensoni Bh. & 

8(7). Anterior pole with epimeritic denticula- 
tions, posterior pole covered by fine 

hairs. Nucleus near one end of the [p. 64. 

body, with single karyosome N. vermicularis Hesse, 

6. Nematocystis hessei Bhatia & Chatterjee. (Fig. 8.) 

"fNematocystis hessei, Bhatia & Chatterjee, 1925, pp. 194-5, pi. viii, 
figs. 11, 11a, 12 ; pi. ix, fig. 24. 
Nematocystis hessei, Cognetti, 1925, p. 231 ; Bhatia, 1929, p. 125. 

Trophozoite elongated like a worm and tapering at either 
end. No epicytal striations. Nucleus long and fusiform, 
with two karyosomes, which are sometimes very unequal in 

Dimensions. — Up to 552 ju, in length by 42 /a in thickness. 



Eemarks. — The species in some respects resembles N. anguil- 
lula, but differs from it in the absence of epicytal striations 
and in the nuclear structure. The nucleus is situated near the 
centre of the parasite and is long and fusiform ; it may 
attain a length of 79 /x. The two karyosomes may be equal 

Fig. 9. 

f '\ 


I I I 

1 ^ ' \ 

Fig. 8. — Nematocystis hessei Bh. & Ch. (After Bhatia and. Chatterjee.) 
Fig. 9. — Nematocystis lumbricoides Hesse. (After Bhatia and Chatter- 
Fig. 10. — Nematocystis pluriharyosomata Bh. & Ch. (After Bhatia and 

or very unequal in size, and are situated near the two ends 
of the nucleus. 

Habitat. — Seminal vesicles of P^.ere^ma Aeferoc^^e to (Mchlsn.) : 
Punjab, Lahore. 


7. Nematocystis lumbricoides Hesse. (Fig. 9.) 

Nematocystis lumbricoides, Hesse, 1909, pp. 153-5, figs. Ixv, Ixvi. 
^Nematocystis lumbricoides, Bhatia & Chatterjee, 1925, p. 196, pi. viii, 

fig. 17. 
Nematocystis lumbricoides, Cognetti, 1925, p. 231 ; Bhatia, 1929, 

p. 124. 

Trophozoite shaped hke an earthworm, swollen in the middle 
and narrower at each end, anterior end somewhat rounded and 
posterior end more pointed. Nucleus long and fusiform, with 
a single karyosome. 

Dimensions. — Up to 1-5 mm. in length and 60 /x in thickness. 

Remarks. — Specimens examined by Bhatia and Chatterjee 
from Pheretima heterochseta (Mchlsn.) at Lahore differed from 
those described by Hesse from Helodrilus caliginosus Savig. 
in France in that the size did not exceed 1 mm. by 31 /x, 
the epicyte was not ornamented with parallel longitudinal 
striations, and the nucleus was variable in position and con- 
tained a single large oval karyosome placed eccentrically. 

Habitat. — Seminsd vesicles o£Pheretim£(, heterochseta (Mchlsn.) : 
Punjab, Lahore. 

8. Nematocystis plurikaryosomata Bhatia & Chatterjee. 
(Fig. 10.) 

■\Nematocystis plurikaryosomata, Bhatia & Chatterjee, 1925, pp. 195-6, 

pi. viii, figs. 13-16 ; pi. ix, fig. 25. 
Nematocystis plurikaryosomata, Cognetti, 1925, p. 231 ; Bhatia, 1929, 
p. 125. 

Trophozoite with a long and extremely deformable body. 
No epicytal striations. Nucleus elongate and oval, with 
several small karyosomes. 

Dimensions. — Up to more than 1 mm. in length and 100/x 
in thickness. Gametocysts small, attaining a size of 140 ^a in 
diameter. Spores 6-5-8-5jU, by 3"4ju, in size. 

Remarks. — The body of the parasite is very deformable and 
shows constrictions and bulgings during the progression of 
the parasite, the granular cytoplasm, together with the 
nucleus, appearing to flow from one pole to the other. The 
parasite differs from N. magna in that it lives free in the 
seminal vesicles, has no polar ornamentations, no hairs round 
its posterior pole, and in its nuclear structure. In stained 
preparations the paraglycogen grains are seen to be aggre- 
gated along the central region, leaving a clear peripheral zone. 
The nucleus is variable in position and contains 6 to 12 large, 
spherical karyosomes ; it measures 60 /x along its long axis. 

Habitat. — Seminal vesicles of Allolobophora (Eisenia) foetida 
(Savig.) : Punjab, KasauH. 



9. Nematoeystis stephensoni Bhatia & Setna. (Fig. 11.) 

■fNematocystis stephensoni, Bhatia & Setna, 1926, pp. 364—7, pis. xi, 
xii, figs. 5-17 ; pi. xiii, figs. 28, 29. 
Nematoeystis stephensoni, Bhatia, 1929, p. 125. 

Trophozoite of an elongate cylindrical form, short and 
swollen or elongated and thin, with blunt extremities. Body 
highly deformable, presenting a number of swellings and 
constrictions. Epicyte shows fine longitudinal striations. 

Fig. 11. — Nematoeystis stephensoni Bh. & Set. 
(After Bhatia and Setna.) 

Nucleus elliptical, with a single central karyosome, which 
consists of a central deeply staining portion surrounded by a 
vacuolated layer. Gametocysts oval. Gametocytes equal in 
size ; gametes similar. 

Dimensions. — Trophozoite up to 1-26 mm. in length and 
100 IX in thickness ; gametocysts from 153 [x by 100 jx to 314 [x 
by 246 IX ; zygotes 8 [x ; sporocysts 15-5 fx by 8-1 fx. 

Eemarks. — In the living condition the parasite is quite 
opaque and capable of contracting and elongating so rapidly 


as to produce constrictions and bulgings, giving it the same 
appearance as N. magna and N. plurikaryosomata. The 
opacity of the granular cytoplasm is so great that the nucleus 
is only faintly indicated. 

In the stained preparations the cytoplasm exhibits the usual 
structure. The nucleus is generally situated about the middle 
of the body, but is often found nearer one or the other extremity ; 
this is due to the movements of the endoplasm. Its elongated 
axis generally Hes parallel to the long axis of the body. The 
shape and structure of the nucleus are very characteristic. 
The nucleus is of an elongate, regularly elUpsoid form, and 
consists of (i) a well-defined nuclear membrane, (ii) a broad 
zone of peripheral achromatin, in which the network is not 
distinguishable, and over which fine chromatin grains are 
dispersed, and (iii) a large central, spherical or oval karyosome, 
which is like an entire nucleus in structure and appears very 
much like a nucleus within a nucleus. The nucleus thus 
appears to consist of two concentric rings and a deeply 
staining homogeneous mass in the centre. The inner ring 
and the central homogeneous mass together constitute the 

Development is on the usual fines, and different stages have 
been described by Bhatia and Setna (1926). 

Habitat. — Seminal vesicles of Eutyphoeus incommodus (Bed- 
dard) : Punjab, Kasauh. 

10. Nematocystis vermicularis Hesse. (Fig. 12.) 

Nematocystis vermicularis, Hesse, 1909, pp. 155-8, figs. Ixvii-lxx 
Berlin, 1924, pp. 68, 74. 
"fNematocystis vermicularis, Bliatia & Chatterjee, 1925, p. 197. 
Nematocystis vermicularis, Bhatia, 1929, p. 124. 

Trophozoite with a fusiform, rather thick body, rounded at 
both extremities. The anterior pole ornamented with a cap 
formed of smaU cylindrical prolongations placed side by side, 
posterior pole covered by fine hairs directed backwards. 
Ectoplasm thin, cuticular ornamentations not visible. Endo- 
plasm rather rich in chromatoid granulations. Nucleus 
eUipsoidal, situated near one of the extremities of the body, 
and containing a single karyosome. 

Dimensions. — Scarcely 1 mm. in length and 100 /x in thickness. 

Remarks. — The single specimen obtained by Bhatia and 
Chatterjee from the seminal vesicles of Pheretima barbadensis 
(Beddard) closely resembled Hesse's fig. Ixviii a of the species 
as descrilDed from Helodrilus longus Ude. The size of the 
specimen was, however, much less, being only 446 /x in length 
by 69jLt in thickness. The nucleus was situated more towards 



the blunt end of the parasite and measured 38-5ju- along its 
long axis. There was a central spherical karyosome. 

Fig. 12. — Nematocystis verinicularis Hesse. (After Hesse.) 

Habitat. — Seminal vesicles of Pheretima barbadensis (Bed- 
dard) : Punjab, Lahore. 

2. Family RHYNCHOCYSTID^ Bhatia, 1930. 

Trophozoites ovoid, spherical or elongated, with a conical 
or cyluidro-conical trunk at the anterior end, soHtary. Sporo- 
cysts biconical, with similar non-appendiculate poles, octozoic. 

Genus RHYNCHOCYSTIS Hesse, 1909. 

Rhynchocystis, Hesse, 1909, p. 45 ; Cognetti, 1911, p. 207 ; Berlin, 
1924, p. 68 ; Cognetti, 1925, p. 232 ; Bhatia & Chatterjee, 1925, 
p. 190 ; Bhatia & Setna, 1926, p. 370 ; Reichenow, 1929, p. 887 ; 
Bhatia, 1929, p. 129; 1930, p. 158; Calkins, 1933, p. 559; 
Troisi, 1933, pp. 327-34. 

Trophozoites ovoid or cyhndroid. Anterior pole provided 
with a metabohc epimerite, which is most frequently elongated 
into a conical or cylindro- conical trunk. Sporocysts with 
characters of the family. 

Key to Indian Species. 

1 (3). Hairs present on both the mucron and 

the posterior end of the body 2. 

2. Form variable, pear-shaped, spherical 
or gregaruiiforni. Epi33ierite meta- 
bolic, a conical or hemispherical mu- 
cron. Nucleus rounded, variable in [p. 67. 
position -B. cognettii Bh. & Ch., 

3 (1). Hairs not present on any part of the 

body 4. 

4 (5). Form elongate, pear-shaped, anterior 

end broader, with a nipple-shaped epi- 
merite. Nucleus oval, usually in [Set., p. 67. 

posterior half of the body E. mamillata Bh. & 




5 (4). Form elongate, cylindrical, with cylin- 
dro-conical epimerite marked with, 
distinct epicytal striations. Nucleus 

oval, usually about the middle of the [p. 66. 

body R. awatii Bh. & Set., 

11. Rhynehocystis awatii Bhatia & Setna. (Fig. 13.) 

fEhynchocystis awatii, Bhatia & Setna, 1926, pp. 371-3, pi. xii, 

figs. 23, 24 ; pi. xv, figs. 34, 35. 
Rhynehocystis awatii, Bhatia, 1929, p. 125. 

Trophozoite with an elongate cyUndrical body, the anterior 
end generally provided with a cyUndro-conical epimerite. 
Epicyte ornamented with fine longitudinal striations, which are 
more distinct and spaced out over the epimeritic region. 
Sarcocyte poorly developed and free from granules. Nucleus 
oval, with a large eccentric karyosome, and generally placed 
about the middle of the body. 

Fig. 13. 

Fig. 15. 

Fig. 13. — Rhynehocystis awatii Bh. & Set. (After Bhatia and Setna.) 
Fig. 14. — Rhynehocystis eognettii Bh. & Ch. (After Bhatia and Chatter- 

Fig. 15. — Rhynehocystis mamillata'Bh.. &c Set. (After Bhatia and Setna.) 


Dimensions. — Maximum length. 400^ ; maximum width, 
in shorter and thicker individuals, 50 jU,. 

Remarks. — This appears to be a rare parasite of its host, 
being found only in a few specimens and in extremely small 
numbers. The form of the trophozoite varies considerably ; 
some are short and thick, while others are long and vermiform. 
An epimerite is not found in all individuals, but, when present, 
it is a distinct club-shaped structure, dilated at its distal 
extremity and narrower at its base. The ectocyte is thin. 
The entocyte is finely granular. The nucleus Hes more often 
in the middle, though in some cases it is found to be situated 
near the anterior end. It is oval in form, with its long axis 
lying along the length of the parasite. The nucleus usually 
measures 17-5 ^u. in its longer and 10-5^ in its shorter diameter, 
and contains a single large eccentrically placed karyosome. 
Some individuals show end to end syzygy ; sometimes the 
satellite retains its epimerite and is fixed by it to the posterior 
end of the primite. 

Habitat. — Seminal vesicles oi Pheretima elongata (E. Perr.) : 
Bombay, Bombay. 

12. Rhynchocystis cognettii Bhatia & Chatterjee. (Fig. 14). 

'fJRhynchocystis cognettii, Bhatia & Chatterjee, 1925, pp. 190—3, pi. vii, 

figs. 1-10 ; pi. ix, figs. 22, 23. 
Rhynchocystis cognettii, Bhatia, 1929, p. 125. 

Trophozoite with an epimerite. Shape of the body variable, 
pear-shaped, spherical or gregariniform, and the anterior end 
provided with a mucron surrounded by a crown of sarcocyte. 
Hairs found only in the region of the mucron and sometimes 
at the posterior end also. Nucleus generally spherical, with 
a single spherical karyosome placed eccentrically and surrounded 
by a clear white halo. The nucleus varies in its position, but 
is never situated in the epimeritic region. Gametocytes are 
nearly equal in size. 

Dimensions. — ^Maximum size of the trophozoite 129 ja by 
46 jU. ; gametocysts are more or less ovoid and measure 
up to 129 ju. by 81 |u,. 

Habitat. — Seminal vesicles of Allolobophora caliginosa 
(Savig.) : Punjab, Kasauli. 

13. Rhynchocystis mamillata Bhatia & Setna. (Fig. 15.) 

^Rhynchocystis mamillata, Bhatia & Setna, 1926, pp. 370—1, pi. xii, 

figs. 21, 22 ; pi. xiv, figs. 32, 33. 
Rhynchocystis mamillata, Bhatia, 1929, p. 125. 

Trophozoite with an elongate pear-shaped body, the 
anterior end broader and provided with an epimerite consisting 
of a nipple-shaped mucron surrounded at its base by a ring 



in which the sarcocyte is well developed. Movements of the 
trophozoite very slow. Body not covered with hair. Nucleus 
oval and situated in the posterior half of the body. 

Dimensions. — The trophozoite may attain a maximum 
length of 126|u,, maximum width 52-5ju, ; nucleus 14-15-9ju, 
in length and S-ll-S/x in width. Cysts and spores not 

Remarks. — R. mamillata differs from R. cognettii in being 
more constant in form and in not exhibiting the changes 
shown by the latter species. It differs from R. pilosa by its 
constant form and by the complete absence of hair covering 
the body. The relative dimensions of the trophozoite, 
nucleus, and karyosome differ markedly from R. awatii, 
which is found in the same host-species, the nucleus being 
larger in trophozoites of corresponding size. 

The sperms of the host, sticking close together to the 
epimerite, form a thick investment round it. The ectoplasm 
is not thick, but the epicyte is fairly weU developed. The 
endoplasm, with the contained granules, is moved about in the 
interior of the body from pole to pole, Avithout affecting the 
external form of the body. The nucleus generally Hes in the 
posterior half of the body, rarely in the middle or the anterior 
half ; it is distinctly oval, and contains a single spherical or 
slightly oval karyosome, which is placed eccentrically. 

Habitat. — Seminal vesicles of Pheretima elongata (E. Parr.) : 
Bombay, Bombay. 

3. FamUy STOMATOPHORID^E Bhatia, 1930. 

Trophozoites ovoid, spherical, cylindrical or cup-shaped, 
"with an anterior sucker-like epimeritic organ ; soHtary. 
Sporocysts navicular, with similar non-appendiculate and 
truncated poles ; octozoic. 

The family includes seven genera (-yic^eBhatia, 1930), of which 
only one is at present known from India. All the genera 
are parasitic in various species of Pheretima only. 

Genus STOMATOPHORA Drzewecki, 1907, emend. Hesse, 
1909, and Bhatia, 1924. 

Stomatophora, Drzewecki, 1907, pp. 218-46 ; Hesse, 1909, pp. 45, 
161-87 ; Bhatia, 1924, pp. 481-512, pi. xxiii, figs. 1-31 ; Bhatia 
& Setna, 1926, pp. 367-8 ; Reichenow, 1929, p. 887 ; Calkins, 
1933, p. 560 ; Ray & Chatterjee, 1936, p. 345. 

Trophozoites ovoid or spherical. Anterior end provided 
with a sucker-like epimeritic organ with or without a central 
mucron. Sporocysts typical of the family. 


Key to Indian Species. 

1 {4). Form with elongate cylindro-conical 

body 2, 

2 (3). Sucker an inverted balloon-shaped de- 

pression with a central mucron and [p. 69. 

radial epicytal striations S. bulbifera Bh. & Set., 

3 (2). Sucker cup -shaped, with a central 

mucron surrounded by a crown of [p. 69. 

petals S. coronata (Hesse), 

4 (1). Form plate- or disc-like, marked by 

furrows and divided into a number of 
irregular lobes. Sucker central, cup- 
like, with a central mucron S. diadema Hesse, p. 71. 

14. Stomatophora bulbifera Bhatia & Setna. (Fig, 16.) 

"f Stotnatophora bulbifera, Bhatia & Setna, 1926, pp. 368-70, pi. xii, 
figs. 18-20, ; pi. xiv, figs. 30, 31. 
Stomatophora bulbifera, Bhatia, 1929, p. 125. 

Trophozoite with an elongate cylindro-conical body, 
a broad and round anterior end and a narrow and pointed 
posterior end. Body not marked by any furrows or epicytal 
striations. The sucker at the anterior end is an inverted 
balloon-shaped depression with a central mucron and radial 
epic3rtal striations. Nucleus, oval with a large spherical 

Dimensions. — Length of trophozoite 56-1 19/x, width 24-5- 
52-5ju. ; nucleus 10-5-15-7/x by 7-11-3/i. 

Remarks. — The greatest width of the organism is at 
one -third the length of the body from the anterior end. 
The sucker is hke an inverted balloon in form, with the epicyte 
Hning it and showing striations diverging from a deeply 
stained central mass, which corresponds to the mucron but 
is not raised. Over the body the epicyte is thm, and the 
sarcocyte forms a thick hyaline and transparent layer. The 
endoplasm is alveolar, and numerous paramylon grains are 
found adhering to the walls of the alveoh. Nucleus with its 
long axis parallel to the long axis of the body. 

Habitat. — Seminal vesicles oi Pheretima elongata (E. Perr.) 
Bombay, Bombay. 

15. Stomatophora coronata (Hesse). (Fig. 17.) 

Monocystis coronatus, Hesse, 1904, p. 268. 

Stomatophora coronata Drzewecki, 1907, pp. 218-46 ; Hesse, 1909, 

pp. 161-87, figs. Ixxiv-lxxix ; pi. ii, figs. 40-73 ; pi. vi, figs. 

136-70 ; pi. vii, figs. 187-93 ; Sokolow, 1911, p. 292 ; Minchin, 

1912, p. 328. 
■\ Stomatophora cwonata, Bhatia, 1924, pp. 499-507, pi. xxiii, figs. 20-5. 
Stomatophora coronata, Cognetti, 1925, p. 233 ; Bhatia, 1929, 

p. 125 ; Reichenow, 1929, p. 887. 

Trophozoite with an oval or elhpsoidal body, rounded at 



both ends. Anterior end bears a sucker, formed as a spherical 
cavity with a cap cut off from the upper pole, and with a central 
mucron arising from the proximal pole as a projection into the 
cavity ; the sucker is surrounded by a crown of a variable 
number of petals formed of hyaline sarcocyte covered over 

Fig. 17. 

Fig. 18. 

Fig. 16. — Stomatophora bulbifera Bh. & Set. (After Bhatia and Setna.) 
Fig. 17. — Stomatophora coronata (Hesse). (After Bhatia.) 
Fig. 18. — Stomatophora diadema Hesse. (After Bhatia.) 

by epicyte marked by striations. Gametocysts generally 
spherical or shghtly elhpsoidal. Sporocysts navicular, but 
their ends are draAvn out and truncated, presenting a button- 
like flattening, sometimes sticking end to end to form chains. 


Dimensions. — Maximum size of trophozoite ISO/a by ISO^u,, 
average 80 ju, by 60 |U, ; gametocysts measure 60-70 ^ by 
50-60 IX ; sporocysts 9-11 ^u by 5-6 jx and 7-7-5 ju. by 3 ft. 

Remarks. — The organism exhibits active though by no means 
rapid movements of the body, thus causing alterations in 
form from moment to moment. The body is generally 
expanded at the anterior end so as to look like a top or a flower- 
vase. It may then show a constriction round the middle of 
the body and assume an hour-glass-like shape, and later a more 
regular outline again. In 1924 I studied the structure of the 
sucker and the crown of petals, and amended the diagnosis 
of the species as given by Hesse (1909). 

There is generally a smaU difference in size between the 
gametocytes enclosed in a cyst, but the gametes do not show 

Habitat. — Seminal vesicles of Pheretima barbadensis (Bed- 
dard) : Punjab, Lahore. 

16. Stomatophora diadema Hesse. (Fig. 18.) 

Stomatophora diadema, Hesse, 1909, pp. 187—9, pi. ii, figs. 66-8. 
f Stomatophora diadema, Bhatia, 1924, pp. 503-7, pi. xxiii, figs. 26—31. 

Stomatophora diademxi, Cognetti, 1925, p. 233 ; Bhatia, 1929, p. 125. 
"f Stomatophora sp., Ray & Chatterjee, 1936, p. 345. 

Trophozoite with the form of a sphere flattened and com- 
pressed between its two poles, so as to resemble a plate or 
disc marked by furrows and divided into a number of irregular 
lobes, some of which are considerably larger than others. 
The sucker is a shallow-like cup depression in the centre of 
the body, and shows a central conical projection or mucron. 
Nucleus generally rounded, sometimes oval, eccentric in posi- 
tion, with a large karyosome. Gametocysts usually spherical. 
Sporocysts navicular, with extremities rather drawn out and 
truncated at the ends. 

Dimensions. — ^Maximum size of trophozoite 105 fx ; gameto- 
cysts 125-170 jjL in diameter ; sporocysts 8-12 /x by 3-4 jx. 

RemarJcs. — In 1924 I discussed the morphology of the 
sucker in S. simplex, S. coronata, and S. diadema, and came 
to the conclusion that, unlike the epimerite of the Polycystid 
Gregarines, the sucker is not present in the earlier stages of 
development in any of the species of Stomatophora, and that 
during the growth of the trophozoite a simple cup-hke sucker 
is formed by a flattening and inpushing of the anterior end of 
the body, the median projection or mucron being thus carried 
to the bottom of the cup-like depression. This process would 
seem to cause the epicytal striations to appear first on the 
surface of the sucker and extending from the central mucron 
to certain definite points on the circular border of the aperture ; 


later they extend beyond over smaller or larger portions of 
the body. In S. simplex these striae do not extend beyond the 
region of the sucker ; in S. coronata they extend over a small 
area of the body immediately surrounding the sucker, thus 
forming a crown of petals ; and lastly, in S. diadema they 
extend not only over the whole of the body, but some of them 
deepen and involve the deeper layers of the ectoplasm, 
causing the body to be cleft into lobes. Ray and Chatter jee 
(1936) have also studied the structure of the suckers of Stomato- 
phora, but their detailed observations have not yet been 

Habitat. — Seminal vesicles of Pheretima barbadensis (Bed- 
dard) : Punjab, Lahore ; seminal vesicles of Pheretima 
posthuma (L. Vaill.) : Bengal, Calcutta. 

4. Family ZYGOCYSTID^ Bhatia, 1930. 

Adult trophozoites always associated in pairs or groups of 
three. Sporocysts biconical, octozoic. 

Genus EXTREMOCYSTIS Setna, 1931. 
Extremocystis, Setna, 1931, pp. 206-9, pi. vi, figs. 2-6. 

Adult trophozoites elongate, resembling Nematocystis ; 
always associated in pairs, attachment being end to end. 

17. Extremocystis dendrostomi Setna. (Fig. 19.) 

■\Extremocystis dendrostomi, Setna, 1931, pp. 206-9, pi. vi, figs. 2-6 ; 
p. 325. 

The only known species in the genus. 

Adult trophozoites have the characters given above ; 
attachment is brought about by one end of an individual 

Fig. 19. — Extremocystis dendrostomi Setna. (After Setna.) 

fitting into a concave depression of the other. Nucleus 
ellipsoidal, greatly elongated, generally lying near that end 
of the trophozoite which joins with the other trophozoite. 


Dimensions. — Maximum size 130 fi by 19 [i, minimum 70 ju. by 

Eemarhs. — The parasites lie coiled and contorted among the 
coelomic corpuscles and the genital products of the host, and 
are never attached to the body- wall or the gut. There is no 
organ of attachment to the host. The body is cylindrical, 
resembhng an elongate Nematode worm, with more or less 
parallel sides and tapering slightly towards the extremities 
of the united pair. Attachment takes place by means of the 
tapering end of an individual fitting into a regular concave 
hemispherical depression of the other. The ratio of width 
to length of each individual is about 1 : 6. Ectoplasm is 
thin. Endoplasm is coarsely granular, and the stream of 
granules flows very swiftly and uniformly from one end of 
the trophozoite to the other, going backwards and forwards 
in a straight line. The nucleus measures 17 /x by Qfi, and Ues 
with its long axis parallel or shghtly inclined to the sides of 
the body. 

The associating pairs become short and pear-shaped and 
later rounded off. Gametocysts not found. Sporocysts 
spindle-shaped, with the two ends finely pointed, 29-35 ju, 
by 6-4 fi. 

Habitat. — Coelomic cavity of Dendrostoma signifer Sel . & de 
Man : Andamans, Port Blair. 

5. Family AIKINETOCYSTIDiE Bhatia, 1930. 

Trophozoites solitary or adherent, branching dichotomously, 
the branches serving for the attachment of the parasite to the 
host. Sporocysts as in Monocystis. 

Genus AIKINETOCYSTIS Gates, 1926. 
Aikinetocystis, Gates, 1926, pp. 400-4 ; Bhatia, 1930, p. 160. 

Trophozoites cyHndrical or columnar, with a characteristic 
regular, dichotomous branching at the attached end, with 
sucker-Hke bodies borne on the ultimate branches ; sohtary 
or in groups of three to eight. Sporocysts as in Monocystis. 
Coelomic parasites in various species of Eutyphoeus. 

18. Aikinetocystis singularis Gates. (Fig. 20.) 

"f Aikinetocystis singularis. Gates, 1926, pp. 400—4, figs. 1—3. 
Aikinetocystis singularis, Bhatia, 1929, p. 126 ; 1930, p. 160. 

Trophozoite cylindrical or columnar, with a characteristic 
regular, dichotomous branching at the attached end ; fixation 



to the host by' means of sucker-like bodies borne on the ultimate 
branches. Ectoplasm shows longitudinal and transverse 
striations. The transparent fluid endoplasm is densely packed 
with ovoid paraglycogen granules. Nucleus is a large ovoid 
body containing a spherical eccentrically situated karyosome. 

Dimeiisions. — Trophozoite may be as long as 4 mm., nucleus 
640-860 IX in length ; gametocysts about 620 ix in length ; 
sporocysts of two sizes, the larger 20-23 yu, long, the smaller 
7-9 /x long. 

Remarks. — The parasite presents a smooth creamy-white 
appearance or, very rarely, a light brownish tinge. The large 
unattached end is bluntly rounded. At a gTeater or less 
distance from this end the body branches into two rami, 

Fig. 20.- 

-Aikinetocystis singularis Gates, n, nucleus ; K, karyosome ; 
Pr, primary ramus ; Sr, secondary ramus. (After Gates.) 

each of which divides to form two smaller secondary rami, 
This branching continues until eight or sixteen small ramuli 
are produced. The dichotomy is regular. The ramuh bear 
groups of irregularly ovoid, sucker-like objects by means of 
which the parasite is attached to the host. Pairs of animals 
are frequently found attached to each other near the rounded 
free ends. Groups of three to eight animals, similarly adherent 
to each other, are also found. 

Living detached specimens show two kinds of movements. 
Waves of peristaltic contraction pass along the trmik and rami 
of the first order, producing a churning of the endoplasmic 
contents in the trunk, from the trunk into the primary rami, 
and back into the trunk ; the nucleus is squeezed to and fro 
from one end of the trunk to the other, or even passing into 


one of the primary rami and returning into the trmik. The 
second kind, described as rotation movements, result in a 
spiral twisting of the primary rami and their branches on their 
own axes. 

Monocystis-hke sporocysts, in masses visible to the naked 
eye, are present in the anterior segment of the host or can be 
found by scraping the body-wall. Ovoid gametocysts were 
found in a few hosts. 

Habitat. — Coelomic cavities of Eutyphceus foveatus (Rosa) 
(most commonly) and E. spinulosus Gates, E. rams Gates, 
and E. peguanus Gates (more rarely) : Burma. 

Genus NELLOCYSTIS Gates, 1933. 
Nellocystis, Gates, 1933, pp. 508-11. 

Trophozoites soUtary or in aggregates of two or three mdi- 
viduals. Anterior end stalk-like and branching into several 
short filaments which attach the parasite to the host. Sporo- 
cysts as in Monocystis. 

19. Nellocystis birmanica Gates. (Fig. 21.) 

Nellocystis birmanica. Gates, 1933, pp. 508-11, figs. 1-8. 

Trophozoites club-shaped, the posterior end enlarged, the 
anterior end narrowing to an elongate stalk. The anterior 
end of the stalk branches into several short filaments, which 
attach the parasite to the host. Within the posterior portion 
is a vacuolar cavity containing a fluid m which there is a smgle 
dark, regularly spheroidal, granular mass contaming a siiagle 
ovoidal nucleus with a single karyosome. Within the stalk, 
near the vacuolar cavity, is a second nucleus, which contains, 
in addition to a large karyosome, a smaller, slightly bent, 
rod-like body. This stalk -nucleus is placed "with its long axis 
along the long axis of the stalk. In some specimens the stalk- 
nucleus is transverse to the long axis of the stalk, and the 
single large karyosome is replaced by a number of smaller 
granules, scattered throughout the nucleus. Sometimes the 
single vacuolar cavity contains two discrete, ovoidal masses 
of unequal size, each containing a nucleus with a single karyo- 
some, and the stalk-nucleus is granular and flattened on the 
side of the vacuolar cavity. Aggregates of two or three uidi- 
viduals, with as many stalks for attachment, are also found, 
and each shows the structure as seen in a sohtary individual. 
Sporocysts are pseudonavi cellar (homopolar), and were seen 
to contain a single nucleus, eccentricaUy located within the 
finely granular contents. 

Remarks. — The Gregarines when present are said to 00010" in 
large number, 50 to 90 per segment for a number of segments. 



The size of the parasites was not determined. The parasites 
were observed in the fluid used for preserving the host, without 
clearing or staining them. In the absence of proper proto- 
zoological technique it is by no means certain that the branching 
stalk was not some tissue of the host, or that the organism 
is really binucleate. Also what is described by the author 
of the species as bi-stalked and tri-stalked forms may, perhaps, 

Fig. 21. — Nellocystis hirmanica Gates, a, portion of a solitary individual ; 

b, portion of the anterior end, more highly magnified ; 

c, bi-stalked form ; d, tri-stalked form. (After Gates.) 

have been syzygies or gametocytes in a process of association 
The organism is so remarkable that it would be well worth 
further detailed study. 

Habitat. — Coelom of the earthworm, Pheretima compta 
Gates : Burma, Toungoo Hills. 

6. Family DIPLOCYSTID^ Bhatia, 1930. 

Trophozoites fusing in pairs to form spherical gametocysts. 
Sporocysts spherical or oval ; octozoic. 

The family includes two genera, Lankesteria and Diplocystis ; 
only the former is known from India so far. 


Genus LANKESTERIA Mingazzini, 1891. 

Lankesteria, Mingazzini, 1891, p. 407, ; 1893, pp. 50, 63 ; Labbe, 
1899, p. 46; Minchin, 1912, pp. 327, 329; Wenyon, 1926, 
pp. 1121-4, fig. 465; Reichenow, 1929, pp. 889-90; Bhatia, 
1930, p. 161 ; Ray, 1933, pp. 392-6 ; Calkins, 1933, p. 559. 

Trophozoites more or less spatulate or leaf-shaped, of small 
size. Epimerite small, described as an anterior pseudopodium- 
like process. Gametocysts spherical, produced by association 
of two individuals after contraction. Sporocysts oval, octozoic. 
Intestinal parasites of Tunicates, Insects, etc. 

Key to Indian Species . 

1 (4). Trophozoite elongate 2. 

2 (3). Trophozoites 50-200 fi. Epimerite 

simple or umbrella-shaped. Nucleus 
spherical, with a single karyosome. 
Gametocysts spherical. Sporocysts 
spindle-shaped L. culicis (Ross), p. 77. 

3 (2). Trophozoite with strongly thickened, 

often spherical anterior end. Nucleus 

large, with one, two, or many karyo- [p. 81. 

somes L.tripteroidesifS^.nov., 

4(1). Adult trophozoite egg-shaped or pear- 
shaped, 101-4 /i by 78 jU. Nucleus 
large, spherical, with a single karyo- 
some. Gametocysts spherical or broad- [Swaminath), p. 79. 
ly oval. Sporocysts spindle-shaped . . L. mackiei (Short & 

20. Lankesteria culicis (Ross). (Fig. 22.). 

^Gregarina culicis, Ross, 1895, p. 346 ; 1898, p. 147 ; 1906, pp. 102, 
Gregarine sp., Marchoux, Salimbeni, and Simond, 1903, p. 713. 
Lankesteria culicis, Wenyon, 1911 a, p. 273 ; 1926, pp. 1121-4, 
fig. 465 ; Knowles, 1928, pp. 495-9 ; Reichenow, 1929, p. 890; 
Lan-Chou, 1930, pp. 361-2, pi. xii, figs. 1-20. 
■ -f Lankesteria culicis, Ray, 1933, pp. 392-6, pi. xxiii. 

Young trophozoites intracellular in the epithelial cells of the 
stomach of the host-larva, developing a simple pseudopodium- 
like epimerite, by which the full-grown trophozoite remains 
attached to the epithelial cell while hanging into the lumen 
of the stomach. Eventually trophozoites become free and 
move about amongst the intestinal contents. The free-living 
forms have a granular cytoplasm, and at the anterior end the 
remains of the organ of fixation, or epimerite, can be detected. 
There is a large central spherical nucleus, with usually a single 
large karyosome. 

When the larva becomes a pupa, the trophozoites leave the 
gut and enter the Malpighian tubes, within which they associate 
in pairs and form large spherical gametocysts. The gametes 
produced by the two Gregarines are of the same size, but 
differ as regards their nuclei ; those produced by one have 

Fig. 22. — Diagram of the life-cycle of Lankesteria culicis (Ross). 
A, escape of eight sporozoites from oocyst, and infection of 
intestinal cells of mosquito larva ; B, growth of young 
Gregarine till it protrudes from the cell though remaining 
attached ; C, free trophozoite in the limaen of intestine ; 
D, association of two sporonts in the Malpighian tube ; 
E-G, associated sporonts in gametocyst, showing nuclear 
multiplication ; H, nuclei arranged on svirface of body ; 
I-J, formation of gametes ; K, fusion of gametes ; L, zygotes 
elongated and encysted in oocysts, in which nuclear multi- 
plication and formation of eight sporozoites is taking place. 
(After Wenyon.) (Xc. 1000.) 


large nuclei and may be regarded as female gametes, while 
those by the other have small nuclei and may be regarded as 
male gametes. The zygotes become elongated and develop 
into spindle-shaped sporocysts ; and in each sporocyst eight 
sporozoites are developed. When the adult mosquito hatches 
from the pupal case, its Malpighian tubes contain gametocysts 
filled with sporocysts. The gametocysts then rupture, sporo- 
cysts escape into the cavity of the Malpighian tubes, and 
make their way to the intestine, whence they pass into water 
to await ingestion by other larvse. 

Dimensions. — A free-hving trophozoite measures from 50- 
200^ in length ; a mature sporocyst measures lOyu, by 6/x. 

Remarks. ~R^y (1933) has pubhshed his observations on 
this form. Accorduig to him two types can be readily 
distinguished. Those which mfest the oesophagus and anterior 
part of the mid-gut, where the chitinous hning is thin and 
smooth, are always of intracellular habit and possess very 
feebly developed epimerites ; while those in the posterior 
part of the mid-gut, where the epithehum is covered by a thick- 
ridged chitin, bear a definite epimerite and are all growing 
extracellularly, only anchored to the epithehum by the 
epimerite embedded m. the host-cell. The epimerite is shaped 
like an open umbrella connected by a very short handle. 
The youngest forms in both cases measure 10/u. by 4-5^. 
At the anterior end of the body there is a clear cone-like zone, 
which takes up a dark, homogeneous stain, and which, according 
to Ray, is comparable with the protomerite of septate Gre- 
garuies. Such differentiation of the anterior portion of the 
cytoplasm is, however, not unknown in other Monoeystids 
possessmg an epunerite {vide Bhatia and Setna, 1927, figs. 18, 
19, & 23), and xmless there is a definite septum the organism 
cannot be placed among the Septata. 

The iatracellular forms, when they become too large to 
remain within the host-cells, enter the gut-lumen, where they 
measure 150-1 94 /z by 3 1-34 /x. The extracellular forms 
attain the same size and, when mature, become detached, 
leaving the epimerite embedded in the epithehum. 

Habitat. — Stomach of the larva, and Malpighian tubes of 
pupa and adult mosquito, Aedes {8tegomyia) segypti (Linn.) : 
India ; and Aedes [Stegomyia) albopictus Skuse : Bengal, 

21. Lankesteria mackiei (Short & Swaminath). (Fig. 23.) 

Certain parasites, Mackie, 1915, p. 948. 

A Gregarine, Christophers, 1924, p. 6 ; Wenyon, 1926, p. 1151. 

Monocystis mackiei. Short & Swaminath, 1927, pp. 539-52, pis. li- 

Uv, figs. 1-18. 
A Gregarine, Knowles, 1928, p. 501. 
Lankesteria mackiei, Reichenow, 1929, p. 890. 



The young intracellular stage occurs in a cell of the intestinal 
epithehum of the larva of a sand-fly (Phlebotomus) , and is at 
first spherical, but later becomes more or less triangular, with 
rounded angles, measurmg about 23-4ju, in length when 
about to leave the host-cell. The adult Gregarine occurs in 
the lumen of the gut or body-cavity of the larva, but only in 
the body- cavity of the pupae and the adult fly. It is egg-shaped 
or pear-shaped, but it can voluntarily lengthen or shorten 
the body and is considerably compressible. Epicyte well 
developed and marked with longitudinal striations. Endo- 
plasm very markedly granular. Nucleus a large spherical 

Fig. 24. 

Fig. 23. 

Fig. 23. — Lankesteria mackei (Short & Swaminath). (After Short & 

Fig. 24. — Lankesteria tripteroidesi, sp. nov. (After Guenther.) 

or subspherical body, measuring about SO fi in its longest 
diameter, eccentrically placed, with a large densely- staining 
spherical or subspherical karyosome, measuring 8-10 /x in dia- 
meter. Gametocj/tes somewhat elongated and crescent- 
shaped, with blunt extremities. They are similar, and a pair 
become apposed and surrounded by a cyst. Gametocysts 
spherical or broadly oval. Hundreds of gametes are formed 
in each gametocyte and are similar in form and size. Zygotes 


secrete an impermeable membrane round themselves and 
become sporocysts. The sporocysts, when released from the 
original gametocyst, are broadly spindle-shaped, terminating in 
a knob-like projection at each end. Each sporocyst contains 
eight sporozoites, apphed closely to one another in a tight 
bundle. Sporocyst ruptures at one or both poles while in the 
ahmentary canal of the larval sand-fly, and the Hberated 
sporozoites attack the epithelial cells of the gut. 

Dimensions. — Young trophozoite (mtracellular) 23-4/x in 
length; adult trophozoite (free in gut) 101-4 /* in length 
by 78 /^ in greatest breadth ; nucleus 30 yu, in length ; karyo- 
some 8-10 /i in diameter ; gametocyst 66-3-105-1 /* in length ; 
sporocyst 9-6 /x by 5-8 /u. 

Remarks. — The complete life -cycle of the Gregarine has been 
described by Short and Swaminath and correlated with the 
different stages in the life-history of the host. The sporocysts^ 
of the gregarine are passed out with the eggs by an adult 
sand-fly. As the larvae hatch out, the sporocysts are taken into 
their ahmentary canal with their first food. The sporozoites^ 
invade the epithelial cells, an intracellular stage is passed, 
and the destruction of the host-cells releases the Gregarinea 
either in the ahmentary canal or the body-cavity of the larva. 
The growth of the larvae is accompanied by the growth of the 
Gregarmes, but in the pupae the Gregarmes in the ahmentary- 
canal have completely disappeared, and in both pupge and 
adults adult Gregarines are only found in the body-cavity, 
where all the other stages of gametogony and sporogony 
take place. 

Habitat. — Ahmentary canal and body- cavity of the larva,, 
and m the body- cavity of the pupa and^ adult of the sand-fly, 
Phlebotomus argentipes Ann. & Brun : Assam ; also in 
P. papatasii Scop., bred out in the laboratory : Bengal,. 

22. Lankesteria tripteroidesi, sp. nov. (Fig. 24.) 

fA Monocystid Gregarine, Guenther, 1914, pp. 264-7, 5 text-figs. 
A Gregarine, Wenyon, 1926, p. 1149. 

Trophozoite elongate, with a strongly thickened anterior 
end which is often spherical and may project laterally. 
Cytoplasm coarsely alveolar and filled with granules. There 
are a few vacuoles at the anterior end. Nucleus large, with 
a shmy protoplasmic thread extending forwards and backwards 
from it. The nucleus contains large dark nucleoli, which 
may be one, two, or many in number. 

Habitat. — Body-cavity, respiratory tubes, and anal gills 
of the larva of the mosquito, Tripteroides dofleini (Guenther), 
{=Ficalbia dofleini (Guenther)) ■ Ceylon. 



2. Tribe HETEROPOLARIDEA Bhatia, 1930 
(including Family Choanosporid^ Dogiel). 

Intestinal or coelomic parasites usually of marine animals, 
such as Polychsetes, Nemertines, Sipunculids, Echinoids, and 
Ascidians. Sporocysts with dissimilar poles. The tribe com- 
prises four famihes {vide Bhatia, 1930), of which only one is 
known, so far, to be represented in India. 

Identification Table of Families. 

1 (2). Sporocysts provided with a typical funnel- [Woodcock. 

like opening at one end Urosporidse * 

2 (1). Sporocysts not provided with a funnel-like 

opening at one end 3. 

3 (6). Sporocysts oval, with a thickening at one 

pole 4. [Kamm, p. 82. 

4 (5). Epimerite simple and deformable Lecudinidae 

5 (4). Sporonts held together by a ball-and-socket [J. S. Huxley. 

joint Ganymedidse * 

6 (3). Sporocysts spindle-shaped, with one side 

slightly more prominent than the other. [Bhatia. 

Gametocysts elongated, sausage-like Allantocystidse * 

Inceet^ sedis. 

Sporocysts not known. Body non-septate, 
with rudimentary epimerite. Develop- [Henry, 

ment intracellular. Sporonts associative . . Kof oidinidse * 

Family LECUDINID^ Kamm, 1922, emend. 
Reichenow, 1929. 

{Syn. DoLiocYSTiD^ Labbe, 1899.) 

Body non-septate, distinguishable from the Monocystids by 
the protoplasm in the anterior portion possessing finer 
granules. A typical epimerite for attachment to the wall of 
the gut of the host is present, but may be lost in fully grown 
individuals. Syzygy does not occur. Sporocysts oval, with 
a thickening at one pole. Parasites of marine Annelids. 

Key to Indian Genera. 

1 (2). Epimerite invaginable, assuming a [p. 83. 

variety of forms Lecudina Mingazzini, 

2(1). Epimerite of a definite form 3. 

3 (4). Epimerite bulb-like at the end of a long [p. 85. 

rigid style Bhatiella Setna, 

4 (3). Epimerite funnel-like, on a long tubular [p. 85. 

stalk Ferbakia Setna, 


Genus LECUDINA Mingazzini, 1891. 

Gregarina (part), Kolliker, 1848, p. 35. 
Lecudina, Mingazzini, 1891, p. 469. 
Doliocystis, Leger, 1893, pp. 204-6. 
Lecudina -\-Ophiodina, Mingazzini, 1893, p. 51. 
Ophioidina, Minehin, 1903, p. 196. 
Doliocystis, MLnchin, 1903, pp. 202-3. 

Lecudina, Kamm, 1922, pp. 10, 23-6 ; Reichenow, 1929, p. 892 ; 
1932, pp. 31-2; 1935, p. 368. 

Body cylindrical or ovoid, cytoplasm of the anterior portion 
distinctly marked ofiF by the possession of fine granules. Epi- 
merite caducous or invaginable, able to take for any single 
species only a variety of well-determined forms. Intestinal 
parasites of Polychaetes. 

23. Lecudina brasili Ganapathy & Aiyar. (Fig. 25.) 
•f Lecudina sp., Ganapathy & Aiyar, 1937, p. 292. 

Youngest stages intracellular within the gut-epithelium, 
ovoid in outline, with a sKght concavity at one end. The full- 
grown trophozoite is widest at about a third of its length 
from the anterior end, and the nucleus, which is spherical, 
is situated at that level. The pellicle is uniformly thick and 
the cytoplasm granular, except in the prolongation at the 
anterior end. These trophozoites lie in the lumen of the gut, 
attached to the epithelial cells by an epimerite. In the 
fuUy evaginated condition the epimerite has a truncated base, 
with a slender prolongation ending ia a darkly staining anchor- 
plate. The peUicle does not appear to be continued over the 
epimerite, which has therefore been interpreted as endoplasmic 
in origin and is capable of retraction and evagination. 
The epimerite is entirely extracellular. In the retracted 
state the anterior region of the organism presents a slight 
concavity. When the organism detaches itself the anchor-plate 
is left behind, and a bubble of cytoplasm appears to protrude 
from the anterior end ; this is in reahty the evaginated epi- 
merite after it has become detached from the gut-epithehum. 
In sections the whole epimerite, excluding the anchor-plate, 
shows faint longitudinal striations. Association takes place 
between two mature trophozoites, which become surrounded 
by a spherical gametocyst. Two kinds of gametes are produced 
and conjugation is anisogamous. Sporocysts are oval in 
outline, with a characteristic thickening at one pole. Eight 
sporozoites are developed inside each spore. 

Dimensions. — Trophozoites measure ISO/x by SOfM, length 
of epimerite 10-12 /x ; gametocysts measure 75 ju, in diameter ; 
sporocysts measure 6 Ju, by 4 |u.. 

Remarks. — Ganapathy and Aiyar (1937) have recently 
communicated a paper describing the different stages in the 




life-cycle of the parasite. Only a brief abstract has beea 
published so far, but through the courtesy of the authors 
I have been able to consult the paper before it appears in print. 
As first shown by Brasil (1908, 1909), there is a well-defined 
" epimerite," but it differs from that organ as it occurs in the 
Cephaline Gregarines. It is an invaginable apparatus, which 
is sometimes like a conical trunk as in Lecudina aphroditse 
(Lankester) or in L. polydorse (Leger) and sometimes like 

Fig. 25. — Lecudina hrasili Ganapathy & Aiyar. Trophozoite attached' 
to the gut-wall. (After Ganapathy and Aiyar.) 

a spherical button as in L. elongata (Mingazzini), but the form 
of the trophozoite is different from any of those species, and 
resembles that of L. pellucida (Kolliker), in which the epimerite 
is described as a simple small papilla. I do not agree with the 
authors in thinking that the epimerite is endoplasmic. Their 
figure shows that the thick pellicle does not extend forward 


■over the epimerite, but they describe striations which are 
certainly epicytal. The so-called epimerite is merely a deform- 
able prolongation of the anterior end, and when the anchor-like 
portion breaks ojff there is a large wound through which the 
entocyte would seem to flow out. The trophozoites are 
parasitized by Metchnikovella sp. 

Habitat. — Intestine of Lumbriconereis sp. : Madras, Madras. 

Genus BHATIELLA Setna, 1931. 
Bhatiella, Setna, 1931, p. 203. 
Epimerite in the form of a bulb-like structure at the end 
of a long rigid style. 

24. Bhatiella morphyssB Setna. (Fig. 26.) 

■\Bhatiella morphysse, Setna, 1931, pp. 203-4, pi. v, figs. 1, 2. 

Solitary, non-septate Gregarine, with a pear-shaped body, 
widest just behmd the middle. Epimerite in the form of 

Fig. 26. — Bhatiella morphysss Setna. (After Setna.) 

a, distinct bulb-like structure at the extreme tip of a long rigid 
style. Dehiscence by simple rupture, no sporoducts. 

Dimensions. — Trophozoite, maximum 200 ^ by 103 /x, 
minimum 100 /x by 40 /i. 

Remarks. — The epimerite is about one-fourth the total length 
of the body, and its style is broad at the base and slender at 
the apex. The protoplasm does not show finer granules in 
the anterior portion of the body. The nucleus is elhpsoidal, 
situated in the posterior half of the body, and contains a 
relatively large karyosome. 

Habitat.— Midi-gut of Morphysa sanguinea Montague i=M. 
Jurcellata Crossland) : Andamans, Port Blau-. 

Genus FERRARIA Setna, 1931. 
Ferraria, Setna, 1931, p. 205. 
Epimerite a wide-mouthed, fimnel-like structure on a long 
tubular stalk. 


Remarks. — Setna (1931) has described the Gregarine as 
septate, and placed it in the family Poljo-habdinidse Kamm. 
Reichenow (1929), however, considers that the presence 
of a septum in Polyrhabdina as described by Kamm is an error, 
and the septate appearance is due to the protoplasm in the 
anterior portion of the body being more finely granular. 
He has consequently amalgamated the family Polyrhabdinidse 
with Lecudinidse. Setna describes and figures the protoplasm 
as dark and finely granular throughout, and indicates the 
septum as a clear area, which may be an artefact. 

25. Ferraria cornucephali Setna. (Fig. 27.) 

■fFerraria cornucephali, Setna, 1931, pp. 205-6, pi. v, figs. 3-5 ; 
pi. vi, fig. 1. 

A soHtary, rather stout-bodied Gregarine. Epimerite a mde- 
mouthed, funnel-like structure on a long, slender, tubular 
stalk. Nucleus large and spherical, with a large qentral 
karyosome. In the digestive tract of a polychsete, quite 
commonly attached to the intestinal wall, or free in the lumen 
of the gut. 

Fig. 27. — Ferraria cornucephali Setna. (After Setna.) 

Dimensions. — Trophozoite, maximum 300 /x by 91 fx, 
minimum 243 jU. by 81 ju. 

Remarks. — Setna has described the form as a septate Grega- 
rme, but this is probably an error (see remarks under the genus). 
He gives ratio of length of protomerite to total length of 
trophozoite as 1 : 4 ; width of protomerite to width of deuto- 
merite as 1 : 1-6. The anterior part (so-called protomerite) 
is hemispherical to subglobular, widest Ijehind. The posterior 
part (so-called cleutomerite) is elongated, cylindrical and 
ovoidal, widest about its middle and well rounded posteriorly. 
There is little or no constriction at the septum, which 
appears to be indicated by a clear area between the anterior 
and the posterior portions. 

Two distinct types of gametocj^sts and sporocysts were 
encountered in the mid-gut of the host, but neither was 
definitely associated with the trophozoites of this or of the 
preceding species. One type consisted of spherical gameto- 
cysts, measuring 90-100 fi in diameter, and full of oval sporo- 
cysts, measuring 10 fx by 4-5 /x. Sporocysts escape by rupture 


of the gametocysts. The other type of gametocyst contamed 
a mass of very pecuUar sporocysts, 20 ju, by 4 ju, ui size, and with 
rounded anterior and posterior ends ; each sporocyst is bi- 
laterally symmetrical, covered by a thick membrane, and 
contains a well-defined nucleus. 

Habitat. — Mid-gut of Morphysa sanguinea Montague (=ilf . 
furcellata Crossland) : Andamans, Port Blair. 

Incert^ sedis. 
Genus DIRHYNCHOCYSTIS Cognetti, 1921. 

Dirhynchocystis, Cognetti, 1921 a, p. 150 ; 1925, pp. 229, 233. 
Echinocystis, Bhatia & Chatterjee, 1925, p. 197. 
Dirhynchocystis, Bhatia, 1929, p. 125 ; Reichenow, 1929, p. 887 ; 
Bhatia, 1930, p. 165. 

Solitary. Body ovoid, with two subcyhndrical appendages, 
arising from opposite sides. Sporocysts biconical, with similar 

Remarks. — Cognetti (1921) describes the two spine-like 
prolongations as anterior and posterior, and regards each as 
comparable to the trunk-hke process arising from the anterior 
end of Ehynchocystis. To me these spines suggest a strong 
comparison with the antero-lateral spines of Ancora, from 
which the form can be derived by shortening and rounding 
off the body. 

26. Dirhynchocystis globosa (Bhatia & Chatterjee). (Fig. 28.) 

"fEchinocystis globosa, Bhatia & Chatterjee, 1925, pp. 197-9, pi. viii, 

figs. 18, 19; pi. ix, figs. 26, 27. 
Dirhynchocystis globosa, Cognetti, 1925, p. 233 ; Bhatia, 1929, 
p. 125 ; Reichenow, 1929, p. 887 ; Bhatia, 1930, p. 165. 

Trophozoite possessing a more or less spherical body, with 
two spine-hke structures radiating from the surface. Nucleus 

Fig. 28. — Dirhynchocystis globosa (Bh. & Ch.). 
(After Bhatia and Chatterjee. ) 

large, generally ovoid, and containing a single karyosome. 
Sporocysts biconical, with two similar truncated poles. 
Dimensions of the trophozoite 74 /a by 65 /x. 


Remarhs. — Infection is very rare. The young trophozoite 
within the blastophore has no spine-like structures. The 
adult trophozoite lives free in the seminal vesicles and possesses 
the spines : it has a dark aspect in the living condition on 
account of reserve of paraglycogen granules, and shows slow 
movements. Form of the adult is like a globe drawn out 
along one axis. The spine-like structures taper gradually 
towards the distal end, and consist of a non-granular endo- 
plasm covered by sarcocyte and epicyte, thus resembling in 
structure the trunk-like epimerite of Rhynchocystis : they 
disappear during association. Nucleus large, slightly elongate 
and oval, generally situated near the middle of the body, 
and may attain a length of 24b fx : it contains a single large 
central karyosome. Sporocysts, doubtfully belonging to this 
species, were unusually large, measuring 28 fi by 14 /x, and 
one of the sporocysts contained only two sporozoites. 

Habitat. — Seminal vesicles of Pheretima posthuma (L. Vaill.) : 
Punjab, Lahore ; Bombay, Bombay. Seminal vesicles of 
Eutyphoeus sp. : United Provinces, Lucknow. 

Genus GRAYALLIA Setna, 1927. 

Grayallia, Setna, 1927, pp. 335-7 ; Bhatia, 1929, p. 126 ; 1930 
p. 165 ; Reichenow, 1929, p. 887. 

Monocystid with four spines at either end. The spine-like 
prolongations arise from the surface of the body at a httle 
distance from the extremities ; they are broader at the base 
than at the apex and are subequal. 

21. Grayallia quadrispina Setna. (Fig. 29.) 

■\Grayallia quadrispina, Setna, 1927, pp. 335-7, figs. 1-3. 

The trophozoite resembles Nematocystis in form, with 
four spine-like processes arising from the surface of the body 

Fig. 29. — Grayallia quadrispina Setna. s, one of the spines. 
(After Setna.) 

at either end and radiating backwards. Body chalky-white 
in colour and highly deformable. Nucleus large and oval, 


containing a variable number of karyosomes. Cysts and 
sporocysts not identified. 

Dimensions. — Size 720-1270 /u, in length by 35-50 /i in width. 

Remarks. — The endoplasm of the parasite is very granular 
and mobile and, owing to its very active movements, the 
parasite assumes a variety of appearances. When extended, 
the breadth is comparatively uniform, but when the parasite 
contracts a characteristic shape is assumed. The epicytal 
striations are well developed over the general body surface ; 
they are faintly visible, and He close together on the spines. 
The sarcocyte is feebly developed and the myocjrte is also 
thin ; both these layers are continued into the spines, but the 
endoplasm is not. The spines are about 12/x in length and 
are capable of movement, being swung actively from side 
to side or held straight against the body. In the endoplasm 
are lodged paraglycogen bodies and many small, oval, and 
sometimes rod-shaped bodies, which may be parasites. 

Habitat. — Seminal vesicles of Pheretima heterochmta (Mchlsn. ) : 
Bombay, Bombay. 

2. Legion SEPTATA Lankester, 1885 

(=Cephalina Delage, excluding DoHocystidae, or Poly- 
CYSTIDEA, sensu stricto). 

The body is divided mto the protomerife and deutomeriie 
by an ectoplasmic septum, and an organ of attachment, 
known as the epimerite, is always present, at least in the earher 
stages. An end-to-end association of two or more trophozoites 
{syzygy) is common : in such the anterior individual is called 
the primite, and the posterior ones the satellites. Parasites of 
the digestive tracts of Invertebrates, especially Arthropods. 

Labbe's classification (1899) into famiHes is generally 
followed with modifications. Kamm (1922) divides the 
group into twelve families. Reichenow (1929) has combined 
two of these, Lecudinidse and Polyrhabdinidse, into one, 
which has been transferred to Haplocyta in this work. 
Reichenow has also transferred the family Porosporidse to 
this group, while two new families, Monoductidse and Hyalo- 
sporinidse, have recently been described by Ray and Chakra- 
varti (1933), and a third, Kofoidinidse, by Henry (1933). 
Of these there does not seem to me to be sufficient justification 
for separating the Hyalosporinidse from the Stenophoridae, and 
Kofoidinidse is placed under Haplocyta in this work. There 
aire thus eleven families in this legion. 


Identification Table of Families. 

1 (8). Sporonts in associations or solitary .... 2. 

2 (7). Sporonts in associations of 2 or 3 3. 

3 (4). Associations of 2 ; form syzygies early. 

Development intracellular. Epimerite 

absent or weakly developed. Gameto- 

eysts dehisce by simple rupture. Sporo- [Kanun. 

cyst ovoidal, with equatorial line .... Cephaloidophoridse * 

4 (3). Associations of 2 or 3. Development 

extracellular 5. 

5 (6). Epimerite a small simple papilla. No 

septum in satellites. Gametocysts de- 
hisce by simple rupture. Sporocysts 

ellipsoidal. Entire life-cycle in a single [L^ger. 

host Didymophyidse * 

6 (5). Epimerite absent or weakly developed. 

Gametocysts dehisce by simple rup- 
ture, setting free " gymnospores." 
Cycle shows alternation of hosts Porosporidse"' Labbe. 

7 (2). Sporonts solitary or in associations up 

to 12. Epimerite simple, symmetrical. 
Development extracellular. Gameto- 
cysts dehisce by simple rupture or by 

spore-ducts. Sporocysts oval or [p. 96. 

barrel-shaped Gregarinidse Labbe, 

8 (1). Sporonts solitary 9. 

9 (10). Epimerite absent or rudimentary. De- 

velopment intracellular. Gametocysts 
dehisce by simple rupture. Sporo- 
cysts ovoidal, with equatorial line, not [Dub., p. 91. 
extruded in chains Stenophoridse Leg. & 

10 (9). Epimerite more or less complex. Devel- 

opment extracellular 11. 

11 (20). Gametocysts dehisce by simple rupture 

or by means of a lateral pseudocyst ... 12. 

12 (17). Gametocysts dehisce by simple rupture. 13. 

13 (14). Epimerite a large cup bordered with 

hooks and placed on a long, slender [Leger. 

neck. Sporocysts crescentic MenosporidSB * 

14(13). Epimerite without a long neck. Sporo- 
cysts crescentic 15. 

15 (16). Sporocysts irregular, biconical or cylin- [Leger, p. 107. 

dro-biconical, not provided with bristles. ActinocephalidSB 

16 (15). Sporocysts with equatorial and polar [Leger. 

spines Acanthosporidse * 

17 (12). Gametocysts dehisce by means of a 

lateral pseudocyst or by simple rupture. 18. 

18 (19). Epimerite asymmetrical, bearing digiti- 

form or root-like prolongations. Sporo- [Leger, p. 108. 

cysts elongate, cylindrical or ellipsoidal. DactylophoridSB 

19 (18). Epimerite symmetrical, with or without 

appendages. Sporocysts pouch-like, [A. Schneider, 

brown or black, in chains Stylorhynchidae * 

20 (11). Gametocysts dehisce by a single elon- 

gated spore-duct 21. 

21. Epimerite a small elevation with prongs 
at its base. Development intracellular. 
Sporocysts compressed, broadly spindle- 
shaped, with median ridge on dorsal [Chak., p. 109. 
surface Monoductid8B Ray & 


1. Family STENOPHORID^ Leger & Duboscq, 


Development intracellular. Trophozoites solitary. Epi- 
merite absent or of a simple structure. Gametocysts dehisce 
by simple rupture. Sporocysts ovoidal, with broad epispore 
and with equatorial line. Not extruded in chains. Parasitic 
in Diplopods. 

Key to Indian Genera. 
1(2). Epimerite absent or rudimentary. Sporo- [p. 91. 

cyst ovoidal, with equatorial line Stenophora Labbe, 

2 (1). Epimerite small, tongue-like, bordered by 

a collar at its base. Sporocyst oval, with [Chak., p. 92. 

a surrounding hyaline membrane Hyalosporina 

Genus STENOPHORA Labbe, 1899. 

Gregarina (part), Frantzius, 1848, pp. 191-4. 

Stenocephalus, Aime Schneider, 1875, p. 584. 

Stenophora, Labbe, 1899, p. 15 ; Minchin, 1903, p. 198 ; Leger & 
Duboscq, 1904, pp. 361-79 ; Watson, 1916, p. 48 ; Kamm, 1922, 
p. 10 ; Reichenow, 1929, p. 894 ; Kudo, 1931, p. 297 ; Calkins, 
1933, p. 561. 

Trophozoites soHtary. Epimerite absent or a mere knob. 
Gametocysts open by simple rupture. Sporocysts oval, with 
a broad epispore and with an equatorial line. Not extruded 
in chains. 

Key to Indian Species. 

Epimerite roimd ; protomerite bottle-shaped, 

with a process at its posterior end; [p. 91. 

deutomerite ellipsoidal. Nucleus elliptical. S. ellipsoidi Chak., 
Epimerite conical or rounded; protomerite 

romided anteriorly and flattened at the 

septum ; deutomerite elongated, tapering [p. 92 . 

posteriorly. Nucleus spherical S. khagendrse Ray, 

28. Stenophora ellipsoidi Chakravarti. (Fig. 30.) 

f Stenophora ellipsoidi, Chakravarti, 1934, pp. 164-8, figs. 1-6. 

Young trophozoite intracellular, with a round epimerite and 
a bottle-shaped protomerite ; not known to penetrate beyond 
the nucleus of the epithehal cells. Sporonts ellipsoidal. 
Epimerite persists for some time. Protomerite in the adult 
has its posterior end drawn out into a small blunt process 
which produces a depression in the septum. Deutomerite 
ellipsoidal, grows more rapidly than the protomerite. Nucleus 
elhptical and variable in position. Gametocysts spherical, 
develop outside the body of the host, and dehisce by simple 
rupture. Sporocysts spindle-shaped, octozoic. 



Dimensions. — Trophozoite, young 8-10-25 ju, in length by 
4/x in width ; sporonts 250-372 /^ by 50-95 /it ; gametocysts 
1 18-172 /x ; sporocysts lO/x by 4/i. 

Habitat. — Mid-gut of a Millipede, Diplopoda sp. : Bengal, 

29. Stenophora khagendrsB Ray. (Fig. 31.) 

■fStenophora khagendrse, Ray, 1933, pp. 343-51, figs. 1-9. 

Young trophozoite intracellular, with a hyaline, conical 
or rounded epimerite. Sporonts solitary, " parrot-shaped." 
Epimerite disappears. Protomerite rounded at the anterior 
end and flattened at the septum, slightly broader than long. 

Fig. 31. 
Fjg. 30. 

Fig. 30. — Stenophora eUipsoida Chak. 

Fig. 31. — Stenophora khagendraa Ray. 

(After Ray.) 

(After Chakravarti.) 
A young tropliozoite. 

Deutomerite up to eight times as long as the protomerite and 
one and a half times as broad, broadest sHghtly behind the 
septum, and in full-grown individuals gracefuUy tapering 
towards the posterior end. Nucleus spherical. Gametocyst 
spherical, dehiscing by simple rupture to release spindle-shaped 

Dimensions. — Trophozoite, young 10-30/x in length by 
6-16/A in width ; sporont, 225/x by 56/x ; gametocyst 100- 
123)U. in diameter ; sporocysts 10-25/x by 4)u,. 

Habitat. — Intestine of a Milhpede (belonging to a new species 
and probably to a new genus) related to Zikadesmus Chamb. : 
Bengal, Calcutta. 

Genus HYALOSPORINA Chakravarti, 1935. 

H yalosporina, Chakravarti, 1935, pp. 211-18 ; Chakravarti & Mitra, 
1936, p. 346. 

Sporonts solitary. Epimerite a small tongue -like elevation 
bordered by a collar at its base. Gametes dissimilar ; during 


fertilization only the nucleus of the male gamete transferred 
to the female gamete. Gametocysts dehisce by simple rupture. 
Spores oval, with a surrounding hyaline membrane. 

Remarks. — Characters such as intracellular development, the 
simple nature of the epimerite, solitary sporonts, and dehiscence 
of cysts by simple rupture clearly indicate the relationship 
of the genus with the Stenophoridae. In the trophozoite the 
nucleus is tethered to the pellicJe by myonemes, in this respect 
resembling Monodttctits, but the spore has a hyaline membrane 
round it and is quite unlike that of either Monoductus or 
Sienophora. On this basis Chakravarti places the genus 
in a new family, Hyalosporinidse, intermediate between 
Stenophoridse and Monoductidae. The genus has so many 
characters in common with the Stenophoridse that I do not 
consider it necessary to place it in a new family on the basis 
of the character of the spore alone, especially when we remember 
that the character of the spore is not known for the majority 
of the species of Stenophora nor for both the other genera 
previously referred to the family. 

Key to Indian Species. 

1(2). Sporonts 800-1 1 1 1 /i by 80-1 1 1 /x. Gam- 

etocysts oval, 292-390 fi by 263-375 fj,. [p. 93. 

Sporocysts oval, 8 /x by 6 jti H. cmnbolopsisie Chalk., 

2 (1). Trophozoites 130-173 /li by 37-70^. 

Gametocysts spherical, 96-120 /x in 

diameter. Sporocysts oval, 6-2 /* by [p. 95. 

4-12 ju H. rayi Chak. & Mitra, 

30. Hyalosporina cambolopsisae Chakravarti. (Fig. 32.) 

^Hyalosporina camholopsisse, Chakravarti, 1935, pp. 211—18, pi. vii, 
figs. 1-19. 

Youngest forms penetrate the epithelial cells, pass beyond 
the nucleus, and develop intracellular ly. Older forms, measuring 
43-1 50 ju- by 14-30 /x, are attached to the epithehal cells by 
an epimerite and grow extracellularly. Epimerite a simple 
structure, consisting of a darkiy staining collar or ring, which 
grasps the host-cell, and a tongue-Uke process, which is 
inserted into the cell. In still older forms very fine root- 
like processes are seen to arise from the ring and project over 
the tongue-like elevation. Protomerite small and conical 
in shape, with its orifice plugged by a darkly staining granule 
which disappears later, the pellicle becoming thickened in 
this region. Deutomerite the longest segment and circular 
in transverse section. Longitudinal epicytal striations more 
prominent on the deutomerite than on the protomerite ; 
a very thin homogeneous layer of sarcocyte, and below that 
a layer of circular myonemes or myocyte to which the nuclear 
myonemes are tethered. Nucleus spherical, measuring 55 /a 



by SBfji, and containing a spherical karyosome measuring 15 /j, 
in diameter. Two sets of myoneme fibres, arising from the 
nuclear membrane, run backwards and are tethered to the 
myocyte in the side-wall. Oval gametocysts are expelled 

Fig. 32. — Hyalosporina camholopsisss Chakravarti. A, yoting tropho- 
zoite growing intracellularly ; B, older trophozoite, with 
epimerite attached by the collar and the tongue-like process 
thrust into the host-cell ; protomerite is at the breaking- 
point, and shows the granule ; C, adult trophozoite, with the 
nucleus tethered to the side-wall by the myonemes ; D, 
mature sporocyst, showing the hyaline membrane, and eight 
_^ ' sporozoites. (After Chakravarti.) 



with the host's excreta and complete development in five or six 
days if kept in a moist chamber. Gametes dissimilar ; male 
gametes have one end pointed and female gametes are spherical. 
During fertiUzation only the nucleus of the male gamete is 
transferred to the female gamete. Gametocysts dehisce by 
simple rupture and release oval sporocysts. A hyaliae mem- 
brane, more prominent at one pole than the other, surrounds 
each spore, and eight sporozoites are arranged superficially 
along the long axis of the spore. 

Dimensions. — Trophozoites, young, attached, 43-150/i by 
14-30 |Lt ; sporonts 800-1111 /t in length by 80-1 llju. in 
breadth ; gametocysts 292-390 ju- by 263-375^ ; sporocysts 
8|U. by 6//.. 

Habitat. — Alimentary canal of a Millipede, Cambolopsis sp. : 
Bengal, Calcutta. 

31. Hyalosporina rayi Chakravarti & Mitra. 

(Fig. 33.) 


■\Hyalosporina rayi, Chakravarti & Mitra, 1936, p. 346 
1936, pp. 116-20, figs. 1-5. 

Sporonts solitary, with early intracellular growt-h. Tropho- 
zoites elongate, the broadest part being slightly behind the 

Fig. 33. — Hyalosporina rayi Chak. & Mit. A, trophozoite; 
B, sporocyst. (After Chakravarti.) 

septum. Epimerite a rounded structure with a darkly 
staining area at its anterior extremity. Protomerite small 


and triangular in shape, with a darkly staining dot or special 
type of granules. Deutomerite broader anteriorly, gradually 
tapering posteriorly, packed with granules. Nucleus spherical, 
with a central karyosome in the young forms, oval in the adult, 
tethered to the pelHcle by two sets of myoneme fibres. Game- 
tocysts spherical, rupturing by simple dehiscence. Sporocysts 
elongate oval, with an outer hyaline coat which is prominent 
at both poles. On one side of the spore there is a circular Hd^ 
like structure which stains deep brown with Lugol's solution. 

Dimensions. — Trophozoites 130-173 /x by 37-70ju. ; gameto- 
cysts 77-116-5 fx in diameter ; sporocysts 6-2 /x by 4-12 fi. 

Habitat. — Alimentary canal of the MiUipedes Polydesmus sp. 
and Strongylosoma contortipes Attems : Bengal, Calcutta. 

2. Family GREGARINID^ Labbe, 1899. 

{Syn. Clepsidrestid^ Leger, 1892.) 

Epimerite symmetrical, simple. Sporonts solitary or in 
associations up to 12. Development extracellular, Gameto- 
cysts with or without sporoducts, sporocysts syrametrical. 
Confined mostly to insects. 

Kamm (1922) enumerates eleven genera as belonging 
to this family, and others have been described since. Only 
five genera are known from India. 

Key to Indian Genera. 

1 (3). Sporonts solitary 2. 

2. Epimerite a simple, sessile knob. Dehis- 
cence by spore-ducts. Sporocysts [p. 97. 
barrel-shaped, in chains Leidyana Watson, 

3 (1). Sporonts in associations of 2 or more 

individuals 4. 

4(7). Sporonts associated in pairs 5. 

5(6). Epimerite a simple globular or cylindrical 
papilla. Dehiscence by spore-ducts. 

Sporocysts barrel-shaped or cylindri- [p. 98. 

cal, in chains Gregabina Dufour, 

6 (5). Satellite with a septum. Epimerite 

dilated anteriorly like a cauliflower 

and narrower at the base. Dehiscence 

by simple rupture. Sporocysts ovoidal [& Set., p. 102. 

or spherical CAtiLOCEPHALUS Bh. 

7 (4). Sporonts in associations of 2 to several 

individuals 8. 

8 (9). Sporonts attenuate, often attached 

laterally. Gametocysts not known. 

Sporocysts barrel-shaped, with a spine [Pinto, p. 103. 

at each corner Pbotomagalh^nsia 

9 (8). Sporonts in associations of 2 to 12 or 

more. EpuTierite a small cylindrical 
papilla. Gametocysts spherical, de- 
hisce by simple rupture. Sporocysts [p. 105. 
ovoidal HiEMOCYSTis Leger, 



Genus LEIDYANA Watson, 1915. 

Leidyana, Watson, 1915, p. 35 ; 1916, p. 44 ; Kamm, 1922, pp. 10, 
72 ; Reichenow, 1929, pp. 895-6 ; Kudo, 1931, pp. 296-7. 

Sporonts solitary, epimerite a simple globular sessile knob. 
Gametocyst possesses spore-ducts. Sporocysts barrel-shaped, 
in chains. 

32. Leidyana gryllorum (Cuenot) Watson. (Fig. 34.) 

Clepsidrina gryllorum, Cuenot, 1897, pp. 52-4. 
Gregarina macrocephala, Labbe, 1899, p. 10. 
Gregarina gryllorum, Cuenot, 1901, pp. 594^5. 
Leidyana gryllorum, Watson, 1916, pp. 120-1, fig. 209. 
^Leidyana gryllorum, Bhatia & Setna, 1924, p. 288. 

Sporonts sohtary, never associative, cylindrical. Ratio of 
length of protomerite to total length 1:5; width of protomerite 

Fig. 35. 

Fig. 34 

Fig. M.~Leidyana gryllorum (Cuenot). (After Cuenot.) 

Fig. ^5.— Leidyana xylocopas Bh. & Set. (After Bhatia and Setna.) 

to width of deutomerite 1 : 1-1. Epimerite globular and 
sessile. Protomerite subspherical, with deep constriction 
at septum. Deutomerite cylindrical, generally conical at the 
end. Nucleus spherical, generally with three karyosomes and 
numerous fine chromatin particles. Gametocysts spherical 
or ovoidal. Spore-ducts 3-8 fx in length ; sporocysts barrel- 

Dimensions. — Sporonts 420 [x m length ; gametocysts 190- 
240 /x in diameter ; sporocysts 7 ju, in longer axis. 



Bemarhs. — Watson (1916) has separated this species from 
L. erratica ; but Bhatia and Setna (1924) are incHned to doubt 
if L. erratica and L. gryllorum are really distinct. Specimens 
having a broadly rounded protomerite may be rounded pos- 
teriorly, and conversely a specimen which might be referred 
to L. gryllorum (by virtue of a deep constriction at the septum 
and a conical posterior end) may possess a protomerite which 
is conical anteriorly. Several specimens show intermediate 
conditions in all these respects. 

Habitat. — Gizzard and mid-gut of Gryllus sp. : Punjab,. 

33. Leidyana xylocopae Bhatia & Setna. (Fig. 35.) 

■\Leidyana xylocopas, Bhatia & Setna, 1924, pp. 279-81, figs. 1-8. 

Body elongate and cylindrical, rounded anteriorly and 
slightly tapering posteriorly, chalky-white in colour and filled 
with paraglycogen granules. Epimerite a large, simple, 
sessile knob, narrowing but slightly near the base. Length of 
protomerite to total length as 1 : 10 ; width of protomerite 
to width of deutomerite as 1:1-4. Sporonts always solitary, 
varying in form and size. Length of protomerite to total 
length as 1 : 4-6-7 ; width of protomerite to width of deuto- 
merite as 1 : 1-3-1-8. Protomerite rounded or dome-shaped, 
and there is a deep constriction at the septum. Deutomerite 
elongate, slightly broader in the middle and tapering 
posteriorly ; posterior end always rounded. Nucleus varies in 
position in the deutomerite, and is a large rounded vesicle with 
a sharply defined membrane and a large spherical karyosome. 
Gametocysts and sporocysts not known. 

Dimensions. — Fairly large cephalont 90 ju. in length; sporonts 
23-174 ^ in length. 

Habitat. — AHmentary canal of the carpenter bee, Xylocopa 
eestuans (Linn.) : Punjab, Lahore. 

Genus GREGARINA Dufour, 1828. 

Gregarina, Dufour, 1828, p. 366. 

Clepsidrina (corr. Glepsydrina), Hammerschmidt, 1838, p. 355, 

Gregarina (part). Stein, 1848, p. 199. 

Gregarina, Schneider, 1873, pp. 515-33. 

Glepsydrina, Aime Schneider, 1875, p. 572. 

Gregarina, Labbe, 1899, p. 9 ; Watson, 1916, p. 43 ; Kamm, 1922, 

p. 11; Wenyon, 1926, p. 1117; Reichenow, 1929, pp. 894-5; 

Kudo, 1931, p. 296 ; Calkins, 1933, p. 561. 

Sporonts associated in pairs. Epimerite a simple globular or 
cylindrical papilla. Gametocysts with spore-ducts ; sporo- 
cysts barrel-shaped or cylindrical, extruded in chains. 



34. Gregarina aciculata, sp. nov. (Fig. 36.) 

fGregarine parasite, "species B," Cornwall, 1915, pp. 130-1, pi. x,. 
fig. 31 ; pi. xi, figs. 37, 38. 

Young trophozoite intracellular in a swollen cell of the 
stomach. Full-grown trophozoite cordate, with a long acicular 
epimerite, penetrating between the epithelial cells right down 
to the basement membrane. Free individuals often assume 
peculiar shapes during syzygy. Sporonts coalesce, acquire 
a common capsule, and become small, white, shining gameto- 
cysts, only just visible with a lens. These cysts are passed out 

Fig. 36. — Gregarina aciculata, sp. nov. A, adult trophozoite ; B, gameto- 
cyst with spores coming out in a long chain. (After Corn- 

with the faeces, and often several are found in the same mass. 
Gametocysts possess a thick, smooth wall and a central sporal 
mass. In the course of seven days or so protrusions from the' 
inside develop into curved or angular tubes about as long as the- 
diameter of the cyst. Sporocysts are then thrust out through 
these tubes in long strings. Sporocysts very small, about 4ju, 
by 2)u,, symmetrically curved, and joined end to end. Dehis- 
cence is along one border. 




Bemarks. — ^This species is more abundant than the larger 
one found in the same host. 

Habitat. — Mid-gut of Lepisma saccMrinalAnn. (?) : Madras, 

35. Gregarina cornwalli, sp. nov. (Fig. 37.) 

fGregarine parasite, " species A," Cornwall, 1915, pp. 128-30, pi. xi, 
figs. 31 A-36. 

Large ; clearly visible to the naked eye when freed from the 
stomach of its host. In the fully developed trophozoite the 
epimerite contains a dehcate transverse septum, and is 
divided off from the protomerite by another well-defined 

Fig. 37. — Gregarina cornwalli, sp. nov. A, adult trophozoite ; 
B, chain of sporocysts. (After Cornwall.) 

septum. A trophozoite does not have its attachment in 
any particular cell ; the globular epimerite appears to be held 
in a cavity, that it has formed, by the pressure of the neigh- 
bouring cells round its neck. Several cells seem to have been 
destroyed to make room for it. In dissections trophozoites 
readily free themselves from their attachments, either leaving 
the epimerite behind or soon casting it off. The remaining 
two segments wander about the field. Deutomerite contains 
an irregular-shaped nucleus in its anterior part. Nucleus 
contains a distinct, centrally placed karyosome, with a ring of 
chromatin material round it. Sporonts coalesce and surround 
themselves with a common capsule. Gametocyst rapidly 
becomes globular and is discharged with the faeces. After about 


twelve days cysts rupture and allow the contained mass of 
sporocysts to escape. The contents of the cyst are long 
strings of sporocysts, interspersed with residual protoplasm. 
Each sporocyst has a thick protective covering and is roughly 
oval, with the curvature greater on one side than the other. 
A sporocyst contains eight shapeless sporozoites, grouped 
four and four at each end. 

Remarks. — The form described above is sufficiently distinct 
from G. lagenoides (Leger), the only species previously known 
from Lepisma saccharina Linn. 

Habitat. — Mid-gut of Lepisma saccharina liinn. (1) : Madras, 

36. Gregarina oviceps Diesing. 

Gregarina achetas abbreviatse, Leidy, 1853, p. 238 ; 1856, p. 47. 

Gregarina oviceps, Diesing, 1859, p. 730. 

Gregarina achetse, Lankester, 1863, p. 94. 

Gregarina achetae abbreviatse, Labbe, 1899, p. 35; Crawley, 1903 a, 

p. 45 ; 1903 b, p. 639; 1907, pp. 220-1 ; EUis, 19136, p. 266 ; 

Watson, 1915, p. 34. 
Gregarina oviceps, Watson, 1916, pp. 101-2, figs. 191-2. 
^Gregarina oviceps, Bhatia & Setna, 1924, pp. 287-8. 

Sporonts bi-associative, obese. Ratio of length of pro- 
tomerite to total length of primite as 1 : 3 or more, width of 
protomerite to width of deutomerite as 1 : 1-1. Protomerite 
hemispherical to subglobose, width twice the length or even 
more. Shght constriction at septum. Deutomerite stout- 
bodied, nearly as wide as long, up to twice as long as wide ; 
widest at shoulder, where it is very little wider than proto- 
merite. Posterior end truncate. Epimerite simple, and 
elongated like a nipple. Endocyte denser in deutomerite than 
in the protomerite, and distinctly marked off from the sarco- 
cyte. Nucleus oval, with fine membrane and many fine 
chromatin granules. Gametocysts spherical. Spore-ducts 
2 to 5, with maximum length of 1000 ^i. Sporocysts barrel- 

Dimensions. — Sporonts, average, 450|u, in length by 225/x 
in width ; maximum length 500 ft ; gametocj'^sts 250 /x in 
average diameter ; sporocysts 4-5 /u, by 2-25 /u.. 

Remarks. — The sporonts examined by Bhatia and Setna 
were shorter and narrower, the ratio of the length of the 
protomerite to the total length of the primite being as 1 : 4 
or even 1 : 5-3 instead of 1:3, as given by Watson. The 
width of protomerite was twice its length or even shghtly 
more. The deutomerite was stout, and varied from nearly 
as wide as long up to twice as long as wide. 

Habitat. — Gizzard and mid-gut of Gryllus sp. : Punjab, 



Genus CAULOCEPHALUS Bhatia & Setna, 1924. 

CaulocepJialus, Bhatia & Setna, 1924, pp. 284-7 ; Ray & Chatter- 
jee, 1936, p. 345. 

Bi-associative ; satelKte with a septum. Epimerite dilated 
anteriorly like a cauliflower and narrowing at the base. Pro- 
tomerite with a characteristic specialized zone anteriorly. 
Oametocysts dehisce by simple rupture. Sporocysts ovoidal or 

37. Caulocephalus crenata Bhatia & Setna. (Fig. 38.) 

■fCaulocephalus crenata, Bhatia & Setna, 1924, pp. 284-7, figs. 12-23 ; 
Ray & Chatterjee, 1936, p. 345. 

Body elongate, cylindrical, densely granular and opaque. 
Epimerite usually dilated anteriorly like a cauliflower and 


Fig. 38. — Caulocephalus crenata Bh. & Set. A, adult trophozoites ; 
B, enlarged view of the epimerite. (After Bhatia and 

narrow basally ; its surface crenate. Protomerite elongate 
and conical and usually 1| times as long as wide, usually 
widest about the middle, with a characteristic specialized 
zone at its anterior end. There is a distinct constriction at 
the septum. Deutomerite cyhndrical, 1| to 3 times as long as 


Ijroad, widest at its middle and gradually narrowing towards 
the rounded posterior end. Nucleus large and spherical or 
somewhat oval, and usually placed about the middle or a little 
in front of it, with a distinct nuclear membrane and one or 
tAvo karyosomes. Length of protomerite to total length as 
1 : 3'3-4*0 ; width of protomerite to width of deutomerite 
as 1 : 1-2-1 -5. In the sporonts the protomerite is consider- 
ably reduced in proportion to the total length of the body. 
Length of protomerite to total length as 1 : 6 ; width of pro- 
tomerite to width of deutomerite as 1:1-6. Sporonts bi- 
associative. Sporonts forming association are much more 
elongate than cephalonts. Gametocysts spherical. Sporo- 
cysts ovoid or nearly spherical. 

Dimensions. — Sporonts 40-142 yu. in total length ; gameto- 
cysts about 90 ju. in diameter ; sporocysts 12 /x. in diameter. 

Remarks. — The epimerite, though presenting a crenated 
surface, cannot be regarded as complex ; it varies in appear- 
£ince and ultimately .becomes simple and symmetrical. The 
protomerite resembles that oiPyxinoides balani and P. cthamali 
{vide Tregouboff, 1912), but the epimerite differs from that of 
Pyxinoides, which is described as a style dilated in the middle. 
Ray and Chatter] ee (1936) have recently studied this parasite 
from the same host at Calcutta, and are of opinion that the 
cauHflower-like appearance of the epimerite is due to the 
action of the fixative or some other disturbing factor. Their 
observations have not yet been pubhshed in full. 

Habitat. — ^Ahmentary canal of the beetle Aulacophora 
foveicollis Kust. i=^A. abdominalis G. et H.) : Punjab, Lahore 
and Kalka ; Bengal, Calcutta. 

Genus PROTOMAGALH^NSIA Pinto, 1918. 

Proto'inagalhaensia, Pinto, 1918 ; 1923, pp. 90-91 (translation, 
pp. 25-6), pis. iii, fig. 48, &vi, fig. 93; Kamm, 1922, p. 11; Setna& 
Bhatia, 1934, pp. 38-42. 

Sporonts in associations of two to several individuals, 
often attached laterally. Sporonts attenuate. Myonemes 
prominent. Epimerite unknoAvn. Gametocysts unknown. 
Sporocysts barrel-shaped, Avith a spine at each corner. 

Remarks. — In addition to the characters given above, 
Pinto (1923) mentions in the diagnosis of the genus that the 
development is always intracellular, and that in the syzygies 
the protomerite of the satelhte embraces the deutomerite of 
the primite, like a pair of forceps. As he places the genus in 
the family Gregarinidae Labbe, in which the development is 
extracellular, the word " intracellular " is probably a mis- 
print for extracellular. 



38. Protomagalhaensia (?) attenuata Setna &;^Bhatia. (Fig. 39.) 

IfProtoniagalhsensia (?) attenuata, Setna & Bhatia, 1934, pp. 40-2, 
figs. 12-22. 

Sporonts attenuate, forming early syzygies. Associations of 

Fig. 39. — Protomagalhsensia (?) attemuxta Set. & Bh. A, free cephalont ; 
B, linear syzygy ; C, lateral attachment of satellites to the 
primite ; D, three satellites attached to a primite ; E, four 
satellites attached to the posterior end of the first satellite ; 
F, two primites followed by a single satellite. (After Setna 
and Bhatia.) 


two to several individuals, often attached laterally. Peculiar 
associations of two primites with a single satelhte sometimes 
met with. Ratio of length of protomerite to total length as 
1 : 13 ; width of protomerite to width of deutomerite as 
1:1-1. Epimerite a simple rounded knob. Protomerite hemi- 
spherical ; deutomerite cylindrical, rounded or flattened 
behind. Satelhte usually longer than the primite, tapering 
posteriorly. Nucleus spherical or oval, with smgle karyosome. 
Gametocysts and sporocysts not known. 

Dimensions. — Associations, maximum length 473 ju,, maxi- 
mum width 22 ju, ; free sporonts 20-70 /x. 

Remarks. — As in Hirmocystis (?) parapeneopsisi Set. & Bh., 
a few cases were seen of a unique type of association in which 
two primites were followed by a single satelhte. Both 
the primites and the satellite were of a slender type and com- 
paratively short in length. 

In the lateral type of syzygy great diversity of form and 
attachment is met with. There may be a satellite in a line 
with the primite, and a second satelhte arise from the side of 
the primite near its posterior end, or both the satellites may 
be attached laterally and diverge like two legs. Sometimes 
there are three satellites, all attached to the posterior end of 
the primite, and in still others several individuals may be seen 
attached to the posterior end of the first satelhte. 

The form resembles Protomagalhiensia in as much as the 
sporonts are attenuate and form associations of several 
individuals, while lateral attachment of sporonts is common. 
Spores in that genus are of a characteristic barrel-shape, with 
a spine at each corner. Until the spores are known the species 
cannot definitely be placed. 

Habitat. — Intestine of the prawn, Parapeneopsis sculptilis 
(Heller) : Bombay, Bombay. 

Genus HIRMOCYSTIS Leger, 1892. 

Eirmocystis (corr. Hirmocystis), Leger, 1892, p. 110. 

Hirmocystis, Labbe, 1899, pp. 12-13 ; Watson, 1916, pp. 167-9 ; 
Kamm, 1922, pp. 11, 74-5 ; Reichenow, 1929, p. 896 ; Calkins, 
1933, p. 562 ; Henry, 1933, pp. 102-9 ; Setna & Bliatia, 1934, 
p. 35. 

Sporonts in associations of two to twelve (or more) in linear 
series, sometimes bifurcated or trifurcated. Epimerite a small 
conical or cylindrical knob, caducous. Gametocysts spherical : 
dehiscence by simple rupture. Sporocysts ovoidal, with two 



39. Hirmocystis (?) parapeneopsisi Setna & Bhatia. (Fig. 40.) 

'\Hirmocystis (?) parapeneopsisi, Setna & Bhatia, 1934, pp. 35-8, 
figs. 1-11. 

Sporonts stoutly built, forming early syzygies. Associations 
of two or three individuals in linear chains, or unique associa- 
tions of two primites with a single satellite. Ratio of length 
of protomerite to total length as 1 : 5-7 ; width of proto- 
merite to width of deutomerite as 1 : 1-4. Epimerite a small 
rounded knob. Protomerite hemispherical or flattened, 
constricted at the septum ; deutomerite cyHndrical or barrel- 
shaped, broadly rounded or flattened behind. Satellite 
longer than the primite, second satelhte longer than the first. 
Nucleus spherical, with a single karyosome. Gametocysts 

Fig. 40. — Hirmocystis (?) parapeneopsisi Set. & Bh. A, early syzygy ; 
B, chain of two individuals ; C, two primites followed by a 
single satellite ; D, single satellite torn asunder by two 
primites pulling apart. (After Setna and Bhatia.) 

spherical or slightly eUiptical. Dehiscence of gametocysts 
and sporocysts not observed. 

Dimensions. — Associations, maximum length 425/x,, maxi- 
mum width 44 /x ; free sporonts 40-80 /x. 

Remarks. — A few cases were met with in this species of 
the rare type of association in which two primites were 
followed by a single satellite. The primites were posteriorly 
rounded off and fitted into curved depressions in the anterior 
margin of the satellite. In other cases two satelhtes ran into 
one another and showed partial or complete fusion, the deuto- 
merite containing one or two nuclei. 

The form resembles Hirmocystis in as much as the sporonts 
form associations of two or more, and the epimerite is small, 


knob-like, and caducous ; but the species cannot be definitely 
placed in that genus until the dehiscence of the cyst and the 
form of the sporocysts has been observed. 

Habitat. — Intestine of the prawn, Parapeneopsis sculptilis 
(Heller) : Bombay, Bombay. 

3. Family ACTING CEPHALID^ Leger, 1892. 

Sporonts always solitary. Epimerite symmetrical, simple 
or with appendages. Gametocysts dehisce by simple rupture. 
Sporocysts irregular, biconical or cylindrical, with conical 
extremities not provided with bristles. 

Genus STEININA Leger & Duboscq, 1904. 

Steinina, Leger & Duboscq, 1904, p. 352 ; Watson, 1916, p. 256 ; 
Kamm, 1922, p. 79 ; Wenyon, 1926, p. 1151 ; Reichenow, 1929, 
p. 900 ; Kudo, 1931, p. 298 ; Calkins, 1933, p. 562. 

Epimerite a short, mobile, digitiform process, changing into 
a flat crenulate disc. Sporocysts biconical. 

40. Steinina metaplaxi Pearse. (Fig. 41.) 

'^Steinina metaplaxi, Pearse, 1933, p. 293, fig. 3. 

Body robust, somewhat flattened. Epimerite nearly spheri- 
cal. Protomerite wider than long, rounded anteriorly. 
Deutomerite rectangular, Avith rounded angles. There is 

Fig. 41. — Steinina metaplaxi Pearse. (After Pearse.) 

a heavy pellicle around the protomerite and the deutomerite. 
Nucleus elHpsoidal, longest axis Ij^ng across body, situated 
usually at end of anterior third of deutomerite, but the position 
varies. Syzygy apparently does not occur. 

Dimensions. — Spdront, length 40-60 /i, breadth 18-20ju, ; 
protomerite 16/i by lOju, ; deutomerite 25ju, by ISjit ; nucleus 
6/a by 7 /A. 


Remarks. — The genus has previously been reported from 
beetles and fleas only. I doubt if the species has been correctly 
referred to the genus Steinina, as in that genus the epimerite 
is a short, mobile, digitiform process changing into a flat 
crenulate disc. 

Habitat. — In the intestine of the Indian crab, Metaplax 
dentipes Heller : Bengal, Port Canning. 

4. Family DACTYLOPHORID^ Leger, 1892. 

Epimerite complex. Sporonts soHtary. Gametocysts dehisce 
with lateral pseudocyst or by simple rupture, Sporocysts 
elongate, cylindrical or elUpsoidal, Parasites in Chilopoda. 

Genus GREENE CKIELL A, nom. nov. 

(=NiNA Grebnecki, 1873). 

Nina, Grebnecki, 1873, p. 264. 

Pterocephalus, Aime Schneider, 1887, pp. 67-8 ; Labbe, 1899, p. 17 ; 

Leger, 1899, pp. 390-3 ; Minchin, 1903, pp. 171, 172, 190, 198 ; 

Leger & Duboscq, 1903, p. 333. 
Nina, Leger & Duboscq, 1909, pp. 33-68 ; Sokolow, 1911, pp. 281-2. 
Pterocephalus, Minchin, 1912, pp. 173, 327, 329, 330, 339. 
Nina, Watson, 1916, pp. 44, 82-4 ; Kamm, 1922, p. 15. 
Pterocephalus, Wenyon, 1926, pp. 74, 1141 ; Knowles, 1928, p. 500. 
Nina, Reichenow, 1929, pp. 904-5 ; Kudo, 1931, pp. 74, 298 ; 

Calkins, 1933, p. 562 ; Reichenow, 1935, p. 370. 

Protomerite formed of two long, narrow, horizontal lobes, 
fused and upturned spirally at one end. Periphery shows 
many teeth, and long slender filaments project from it. 
Cyst dehisces by pseudocyst. Spores long-ovoidal, united 
in oblique chains. In the intestine of Mjrriopods. 

Remarks. — The name Nina is preoccupied for a moUuscan 
genus, Nina J. E. Gray, 1850. The second nsbiae, Pterocephalus, 
is also inadmissible, being preoccupied for an elasmobranch 
fish, Pterocephalus Swainson, 1838. It may incidentally be 
remarked that the name Pterocephalus has also been wrongly 
used for a Nematode (0. Linstow, 1899) and a Trilobite 
(F. Raw, 1907). I have, therefore, renamed the genus 
Grebneckiella, after the author of the type-species. 

41. Grebneckiella navillae (Mitra & Chakravarty). 

■^Nina navillse, Mitra & Chakravarty, 1937, p. 1. 

Sporonts soHtary. Gametocysts oval. Sporocysts spherical 
to oval, with two envelopes ; liberated in chains. 


Dimensions. — Sporonts 819-975 ju, by 97-190 /x ; gametocysts 
125-175^ by 95-125^. 

Eemarks. — Full observations have not yet been published, 
and it is not certain that the form is distinct from other 
previously known species. 

Habitat. — In the intestine of Scolopendra sp. : Bengal, 

5. Family MONODUCTID^ Ray & Chakravarty, 


Sporonts solitary. Epimerite a small elevation with prongs 
attached to its base. Gametes dissimilar. Gametocysts 
dehiscing by a single elongated spore-duct. Sporocysts 
compressed, of broad spindle-shape, with median ridge on 
dorsal surface. 

Genus MONODUCTUS Ray & Chakravarty, 1933. 
Monoducttis, Ray & Chakravarty, 1933, p. 359. 
With the characters of the family, 

42. Monoductus lunatus Ray & Chakravarty. (Fig. 42.) 

■\Monoductus lunatus, Ray & Chakravarty, 1933, pp. 352-60, pi. ii, 
text-figs. 1-5. 

Young trophozoite intracellular. Full-grown trophozoite 
elongate, widest shghtly behind the septum. Epimerite 
knob-like, with twelve to sixteen stiff, radiating processes 
attached to its neck. Protomerite more or less conical, 
relatively very small, with an orifice at its anterior extremity, 
through which longitudmal myonemes run connecting the 
base of the epimerite with a raised disc-like platform arising 
from the middle of the septum. There is a series of transverse 
myonemes also, just underlying the pelhcle. Deutomerite 
elongate, circular in transverse section, with circular myonemes 
well developed, and with the cytoplasm usually packed with 
granules, which give the organism an opaque yellowish-white 
appearance. Nucleus contains a single spherical karyosome, 
and is shaped like a parachute, a number of fibrils running 
back from its posterior concave surface which vary the form 
and position of the nucleus. Prior to association, sporonts 
develop posterior pseudopodial processes hj which couphng 
is effected. A thin transparent cyst is formed round the paired 
gametoc3Ates. The spherical gametocysts are expelled with 
the host's excreta and complete their development in three 



or four days if kept moist. There is no distinction between 
the gametocytes, but one gives rise to slightly drawn out 
and pointed (male) gametes and the other to perfectly 
elliptical (female) gametes. Sporocysts compressed, truncate 
at one pole, and with a median ridge. They are released 
through a single spore-duct in a long chain formed of pairs 
set obliquely. 

Fig. 42. — Monoductus lunatus Ray & Chak. A, adult; B, same, 
showing the orifice in the protometerite and the longi- 
tudinal myonemes passing through it. (After Ray and Chak- 

Dimensions. — Full-grown trophozoite 225-445 /z. by 33- 
47 /x ; length of epimerite 7-5-10-25/i; nucleus 30/i by 12ju,; 
gametocysts 225-230 ju, ; sporocysts 10-25ju, by 4jU,. 

Habitat. — Alimentary canal of a Millipede, Strongylosoma 
contortipes Attems : Bengal, Calcutta. 




Leger, 1900. 

(Syn. Amgebosporidia A. Schneider, 1884.) 

The Schizogregarines are parasites of the digestive tract and 
appended organs (e. g., Malpighian tubules) of Arthropods, 
AimeUds, and Tunicates. The sporocyst gains entrance into 
the digestive tract of the specific host and the sporozoites are 
set free. These develop into trophozoites either in the lumen 
of the gut or within the cells of the host and undergo schizogony. 

Fig. 43. — Life-cycle of a typical Schizogiegarine, Schizocystis gregari- 
noides Leger. A, sporozoite escaping from the spore ; B-E, 
growth of the sporozoite into the multinucleate schizont, of 
which there are two types, the vermiform schizont (a) which 
attaches itself to the epithelial cell by its anterior end, and 
the massive schizont (6) which lies free in the gut of the host ; 
F, division of the schizont into a munber of merozoites, which 
may grow into schizonts again (G^, G^), or grow into gamonts 
((?*) ; H, yoiuig gamonts ; /, association of two full-grown 
garaonts ; J, formation of a common cyst round the two ; 
K, division of nuclei in each gametoeyte ; L, formation of 
gametes ; M, fusion of gametes in pairs ; N, each zygote 
becomes an oocyst and develops eight sporozoites in it. 
(After Leger.) 


which may be binary or multiple fission or budding. As in 
Ettgregabinakia, two full-grown trophozoites (sporonts) 
associate and give rise to gametes, which conjugate to produce 
zygotes. Each zygote becomes an oocyst. The number of 
oocysts produced by each pair of gametes varies from one to 
thirty. Each oocyst, popularly called a spore, contains one 
to eight sporozoites. 

L6ger and Duboscq (1908) divided the suborder into Poly- 
sporidea and Monosporidea according to the number of 
oocysts produced by a pair of associated Gregarines. Fantham 
(1908) on the other hand divided the group into Endoschiza 
and Ectoschiza, according as schizogony took place within a 
cell or was extracellular respectively. Keilin (1923) is of the 
opinion that neither of these classifications is a natural one, 
and that it is not impossible that the process of schizogony 
has been secondarily acquired by some members of the 
EuGREGARiNARiA and that the various genera of Schizogre- 
GARINARIA wiU ultimately be distributed among the famihes 
of the EuGREGARiNARiA. Wenyon (1926) and Ray (1930) 
have supported this view. 

Reichenow (1929) does not group the genera into families, 
but Minchin (1912), Doflein (1916), and Calkins (1926) recog- 
nized a number of families, each based on one or two genera. 

Identification Table of Families. 

1 (7). Gametocyst contains a single oocyst . 2. 

2 (4). Oocyst contains a single sporozoite . . 3. 

3. Gregarine spirally wound or cres- [Dub. 

centic ; schizogony intracellular . . . Spirocystidse * Leg. & 

4 (2). Oocyst contains eight sporozoites ... . 5. 

5 (6). Gregarine ovoid, with an epimerite ; [Porter. 

schizogony intracellular Merogregarinidse * 

6 (5). Gregarine conical, with pseudopodial 

processes along its base ; schizogony [& Dub. 

extracellular Ophryocystidae * Leg. 

7 (1). Gametocyst contains many oocysts . . 8. 

8 (15). Only one type of schizogony 9. 

9 (12). Schizogony intracellular 10. 

10 (11). Gregarine elongated, with longitudinal 

striations, and chromatic bodies at 
the anterior end. Gametocyst con- 
tains many oocysts, each containing 
four or eight sporozoites Selenidiidae * Brazil t. 

11 (10). Gregarine elongate. Gametocyst con- 

tains sixteen oocysts, each containing 

eight sporozoites Lipotrophidae * Keilin. 

12 (9). Schizogony extracellular 13. 

13 (14). Gregarine vermicular. Gametocyst 

contains large number (about thirty) 

of oocysts, each containing eight [Dub. 

sporozoites Schizocystidse * Leg. & 

■)• According to Ray (1930) certain species of Selenidium do not 
show any schizogony. 


14 (13). Gregariiie ovoid, with a process for 

attachment. Gametocyst contains 
eight oocysts, each containing eight 
sporozoites CauUeryellidse * Keilin. 

15 (8). Two types of schizogony, one intra- 

cellular and the other extracellular. 

Gametocyst contains a large number 

of oocysts, each containing eight [yavlensky. 

sporozoites Menzebieridse * Bogo- 

Up to the present time nobody has worked on the 
Schizogregarines of India. 

II. Order COCCIDIA Leuckart, 1879. 

The CocciDiA have a wide distribution, occurring as para- 
sites in the Vertebrates as well as the higher Invertebrates. 
They are generally found as parasites of the epithelium of the 
digestive tract and associated glands. Alternation of genera- 
tions is invariably present : asexual reproduction is by 
schizogony, and is followed by sexual reproduction by, in 
most cases, anisogamy. During the whole of their growth 
the male and the female gametocytes are apart and develop 
independently of one another. The male gametocyte generally 
produces a relatively large number (six or more) of male 
gametes. Both kinds of reproduction take place in the body 
of one and the same host (except in the Aggregatidse and 
the Hsemogregarinidse). The Coccidia are divided into two 
suborders, as follows : — 

1. Gametocytes dissimilar in size ; associated 

with each other during the later part of [p. 113. 

trophic life ; microgametes few Adeleidea Leger, 

2. Gametocytes similar in size ; independent, 

each microgametocyte developing into [p. 156. 

numerous microgametes Eimeridea Leger, 

I. Suborder ADELEIDEA Leger, 1911. 

The scbizonts develop into micro- and macrogametocytes 
which become closely associated and develop in contact with 
one another. The microgametocyte produces a few (two or 
four) microgametes. The zygote divides into numerous 
sporoblasts, each of which develops into a sporocyst with two 
or four sporozoites. These sporozoites, which are small 
gregarinulse, penetrate the epithehal cells of the host and 
grow into large rounded or oval schizonts (agamonts). The 




nucleus undergoes repeated division and the schizont divides 
into a corresponding number of merozoites, which are set 
free within the lumen of the organ. Each merozoite infects 
another cell. After several such generations the merozoites 
develop within the cell into micro- and macrogametocytes. 

Leger (1911) divided the suborder into three famihes, 
Hsemogregarinidse, LegerelUdae, and Adeleidse, on the basis 
of the number of sporocysts and contained sporozoites. 
Reichenow (1921 and 1929) grouped all the genera in a single 
family, Adeleidae Leger, emend. Noller (1928), after con- 
sideration of the various schemes proposed, is in favour of 

Fig. 44. — Stages in the life-cycle of Adelina dimidata, a typical member 
of the suborder. ( x 1700.) A, association of macroga- 
metocyte and microgametocyte ; B, nuclear division in 
microgametocyte and formation of gametic nuclei ; C, sporo- 
cyst with two sporozoites. (From Wenyon, after Schellack.) 

employing Leger's system, as a matter of practical con- 
venience, until a generally recognized natural system can 
be adopted. 

Followiag Kudo (1931), the suborder is here divided into 
two families, Adeleidse and Hsemogregarinidse, which corre- 
spond to the two suborders of similar name of Wenyon 

ADELEA. 115 

Identification Table of Families. 

1 (2). Parasitic in the epithelium of the digestive 

tract and its appended glands, chiefly of 

Invertebrates. Zygote motionless ; be- 
comes enclosed in a resistant oocyst, 

which does not increase in size. The 

sexual and asexual cycles occur in one [emend., p. 115 

host Adeleidae Leger, 

2(1). Parasitic in the cells of the circulatory 

system of Vertebrates. Zygote motile ; 

forms an oocyst which increases in size. 

Alternation of hosts, asexual cycle in 

a Vertebrate and sexual cycle in an [Leger, p. 117. 

Invertebrate Haemogregarinidse 

1, Family ADELEIDiE Leger, 1911, emend. 

Two gametocytes, one of which is smaller in size, unite in 
a kind of pseudo-conjugation. The nucleus of the micro- 
gametocyte divides once or twice, and one of its products 
enters the macrogamete and fuses with its nucleus. The 
zygote is motionless and becomes enclosed in a resistant 
oocyst, which does not increase in size. The sexual and 
asexual cycles occur in the same host. They are intestinal 
parasites of Invertebrates, and infection is contaminative. 

Wenyon (1926) raised the group to the rank of a suborder, 
under the title Adblejdea, and divided it into four famihes, 
viz., DobelHdse Ikeda, 1914 ; Legerellidse Leger, 1911 ; 
Adeleidae Mesnil, 1903 ; and Klossielhdse Wenyon, 1926. 
Reichenow (1929) and Kudo (1931) place the family Dobellidae 
in the suborder Eimeridea. The other genera so far known 
can conveniently be treated as belonging to a single family. 

Key to Indian Genera. 

1 (2). Oocyst thin-walled, containing numerous [p. 115. 

disc-shaped spores Adelea Schneider, 

2 (1). Oocyst thick-walled, containing smaller [p. 116. 

number of spherical spores Adelina Hesse, 

Genus ADELEA Aime Schneider, 1875. 

Adelea, Aime Schneider, 1875, p. 598 ; Schaudiim& Siedlecki, 1897, 
p. 193 ; Labbe, 1899, p. 56 ; Perez, 1899, pp. 694-6 ; Minchin, 
1903, p. 233 ; Dobell, 1907, pp. 155-62 ; JoUos, 1909, pp. 249- 
62 ; Schellack & Reichenow, 1910, pp. 380-3 ; 1915, pp. 425-60 ; 
Leger, 1911, pp. 71-88 ; Debaisieux, 1911, pp. 259-66 ; Minchin, 
1912, p. 352 ; Greiner, 1918, pp. 522-9 ; Wenyon, 1926, p. 1069 ; 
NoUer, 1928, p. 189 ; Reichenow, 1929, p. 912 ; Kudo, 1931, 
pp. 277-8 ; Calkins, 1933, p. 544. 

The zygote develops into a large thin-walled oocyst which 



contains a variable number of flattened sporocysts, each with 
two sporozoites. Other characters as in the suborder and the 

43. Adelea pachelabrse de Mello. 

■fAdelea pachelabrse, de Mello, 1921, p. 242. 
Adelea pachylabree, Wenyon, 1926, p. 1069. 
Adelea pachelabrsR, Reichenow, 1929, p. 912. 

Schizonts show a sexual dimorphism. Macroschizonts 
rounded or oval, with the cytoplasm markedly alveolar, 
containing granules of reserve material, and the nucleus 
surrounded by a distinct membrane. Binary fission equal or 
unequal, leading to formation of up to sixteen nuclei. Micro- 
schizont has a feebly alveolar, non-granular cytoplasm, 
and the nucleus is compact and stains deeply. Macro- 
merozoite is oval, with alveolar cytoplasm and spherical 
nucleus. Micromerozoite is oval or fusiform, and its nucleus 
contains a distinct karyosome. Fertihzation is preceded by 
the association of gametes, one or several male gametes 
(micromerozoites) being attached to a female gamete (macro- 
merozoite). Female gamete is rounded oval, and possesses 
a voluminous nucleus which does not show a karyosome. 
Male gamete is generally smaller and attaches itself to one 
of the poles of the female gamete. Zygote develops into an 
oocyst with two binucleate sporoblasts, each nucleus repre- 
senting a future sporozoite. Sporocysts dizoic. 

Remarks. — Wenyon (1926) considers it doubtful if this 
parasite belongs to this genus. 

Habitat. — Intestine and digestive glands of a Mollusc, 
Tachelahra mosstra Reeve : Porttjguesb India, Nova Goa. 

Genus ADELINA Hesse, 1911. 

Klossia (part), Aime Schneider, 1885, p. 7. 

Adelea (part), Labbe, 1896, p. 536 ; 1899, p. 56 ; Minchin, 1903, 

pp. 233, 235, 332. 
Adelina, Hesse, 1911, pp. xv-xix ; Wenyon, 1926, pp. 1070-2; 

Reichenow, 1929, pp. 912-4 ; Kudo, 1931, p. 278 ; Calkins, 1933, 

p. 566 ; Reichenow, 1935, pp. 371-2. 

Oocyst thick-walled. Sporocysts spherical and compara- 
tively few in number. 

44. Adelina schellacki Ray & Das-Gupta. 

Adelina schellacki, Ray & Das-Gupta, 1937, p. 291. 

Oocyst oval or egg-shaped, without residue ; eight spherical 
sporoblasts, with two sporozoites, developed in each within 
10 to 15 days if the unsegmented oocysts are kept in a moist 
chamber. The oocystal membrane ruptures after being in 


the moist chamber for four or five days, and along with this 
the unused microgametes are discarded. 

Habitat. — Intestine of the centipede, Cormocephalus dentipes 
Poc. : Bengal, Calcutta. 

2. Family H^MOGREGARINID^E Leger, 1911. 

The researches of Reichenow and others have shown the 
coccidial nature of the Hsemogregarines, long known to occur 
in the red or white blood-corpuscles of all classes of Vertebrates. 
They are in reahty Coccidia, which have certain stages adapted 
to life within the circulating cells of the Vertebrate blood. 
Schizogony may or may not take place in the blood- corpuscles, 
but the gametocytes always enter the blood- corpuscles. The 
fertilization process is of the Adelea-type. The formation of 
oocyst and sporozoites takes place in the body of the blood- 
sucking Invertebrate, which in turn transfers the sporozoites 
to a second Vertebrate host. 

The group is also related to Lankesterelhdse, in which the 
life-cycle is typical of that of Eimeridea and takes place either 
in the intestine (the usual habitat of Coccidia) or in endothehal 
cells of the blood-vessels. 

Wenyon (1926) raised the group to the rank of a suborder, 
under the title Haemogregarinidea, and divided it into three 
families, viz., Hsemogregarinidse Neveu-Lemaire, 1901 ; Hepa- 
tozoidse Wenyon, 1926 ; and Karyohsidse, Wenyon, 1926. 
Reichenow (1929) included the Hsemogregarines, as well as 
Adelea and related genera, in a single family, Adeleidse Leger 
emend. I have, however, followed Kudo (1931) m placing 
the Hsemogregarines in a separate family from the Adeleidse. 

Key to Indian Genera. 

1 (3). Schizogony in the red. blood-corpiiscles 

of the internal organs of a Vertebrate . 2. 

2. Gametogony and fertihzation in the 
body of blood-sucking Invertebrate 
(leech). Oocyst small, directly pro- 
ducing eight sporozoites, without form- [Danilewsky, p. 119> 
ing sporocysts H^MOGBEGAnnsrA 

3 (1). Schizogony not in the red. blood-cor- 

puscles 4. 

4 (5). Schizogony in cells of the internal organs 

(liver, spleen, bone-narrow, etc.) or 
leucocytes of Vertebrates. Gameto- 
gony and fertilization in the body of 
a blood-sucking Invertebrate (tick, 
mite, etc.). Oocyst increases enor- 
mously in size, producing sporoblasts, 

sporocysts, and sporozoites in the [p. 145. 

oocyst Hepatozoon Miller, 

Fig. 45. — Life-cycle of Hxmogregarina stepanowi Danilewsky. The 
figures to the right of the dotted Une represent the phases 
in the blood of the tortoise ; those to the left the phases in 
the leech. A, sporozoite ; B, C, early schizogony producing 
a large nuinber of merozoites ; D, merozoite penetrating a 
blood-corpuscle ; E-H, later schizogony, in which few 
merozoites are produced ; in i'^ is seen the recurved vermicule 
within the corpuscle ; /, free merozoite about to enter 
a corpuscle and repeat the stages D-H, or to initiate the next 
phase ; J, K, final schizogonous generation which produces 
the gametocytes ; L^, Lc^, sexually differentiated merozoites 
which grow up into micro- (Mj) and macrogametocytes {M^) 
respectively, and develop further in the leech when taken 
up by it ; N, association of micro- and macrogametocytes 
in the gut of the leech ; O, formation of four microgametes 
by the microgametocyte ; P, one of the microgametes has 
penetrated the body of the macrogamete and fused with its 
nucleus ; Q, zygote with synkaryon, the degenerating 
remains of the male gametocyte are attached to it, and are 
also seen in the next four stages ; R, S, T, successive divisions 
of the synkaryon ; U, ripe cyst containing eight sporozoites. 
(From Minchin, after Reichenow.) 



5 (4). Schizogony in the endothelial cells of 
blood-vessels of a Vertebrate. Game- 
togony and fertilization in the body of 
a mite. Oocyst produces sporoblasts, 
which are liberated as motile vermicules 
and infect the host's eggs. The mite, 
hatched from the egg, has the sporo- 
cysts in its intestinal epithelium. These 
are cast off and voided with the faeces, 
which are eaten by the Vertebrate host. 

The sporozoites make their way to the [p- 154. 

endothelial cells Kaeyolysus Labbe, 

Genus HiEMOGREGARINA Danilewsky, 1885. 

Hsemogregarina, Danilewsky, 1885, pp. 588-98. 

Danilewskya, Labbe, 1894, p. 124. 

Danilewskya+Laverania, Billet, 1895, p. 30. 

Hsemogregarina, Labbe, 1899, pp. 76-7 ; Minchin, 1903, pp. 265, 

266-7, fig. 77 ; 1912, pp. 372-6 ; Wenyon, 1926, pp. 1081-4 ; 

Reichenow, 1929, pp. 924-7 ; Thomas & Robertson, 1929, 

pp. 104-5, 109-10 ; Kudo, 1931, pp. 279-81 ; Reichenow, 1932, 

p. 45 ; Calkins, 1933, pp. 545, 567. 

Schizogony takes place in the red blood- corpuscles or other 
cells of the body of Vertebrates, merozoites escape from 
the original host-cell and infect other corpuscles or cells. After 
several generations merozoites develop into gametocytes in 
the red blood- corpuscles. These are taken up with the blood 
by an Invertebrate (leech), in the gut of which association 
of the micro- and macrogametocytes occurs. The gameto- 
cytes escape from their host-cells, become more or less spherical, 
associate in pairs, and a cyst-wall is formed, enclosing both the 
macro- and microgametocyte. The microgametocyte produces 
two to four microgametes, and the macrogametocyte forms 
a single macrogamete. One of the microgametes fertihzes the 
adjacent macrogamete, and a small oocyst is formed. The 
oocyst directly produces eight sporozoites without the formation 
of sporocysts. 

The genus was founded by Danilewsky for the Hsemogre- 
garine H. stepanowi Danilewsky of the European tortoise. 
The life-history was described by Reichenow (1910) and may 
be followed from fig. 45. 

45. Hsemogregarina berestneffi Castellani & Willey. (Fig. 46.) 

^Hsemogregarina sp. (probably new), Berestneff, 1903, pp. 343-8, 

pi. viii, figs. 1-9. 
Hsemogregarina berestneffi, Castellani & Willey, 1905, p. 397. 
"^Hsemogregarina sp., Patton, 1908, p. 319. _ 

\Hsemogregarina berestneffi, Dobell, 1910, p. 67, pi. u, figs. 3-8._ 
Hsemogregarina berestnieffi, de Mello, de Sa, de Sousa, Dias, & 

Noronha, 1917, p. 13. 
"^Hsemogregarina sp., Donovan (first recorded m Wenyon, iy/0, 

Hsemogregarina berestneffi, Wenyon, 1926, p. 1398 ; Scott, 1926, 

p. 238. 
Hsemogregarina encapsulse, Wenyon, 1926, p. 1398. 


Intra-corpuscular individuals of various forms and sizes, 
many of them showing a characteristic pink-staining sheath. 
Free gregariniform individuals actively motile in the blood 
plasma. Small forms enter by boring directly into the cor- 
puscle. Occasionally the organism, on reaching the inside 
of the corpuscle, rests for a few minutes and then wriggles out 
again into the plasma. 

Dimensions. — Length 26-28 /u, ; mdth 4-5- 5 /x. 

Fig. 46. — Heemogregarina berestneffi Cast. & Will. A, intra-corpuscular 
form ; B, free form. (After Dobell.) 

Remarhs. — ^According to Berestneff the parasite lies within 
a strongly refringent colourless capsule, both ends of which 
are rounded and curved, the whole embracing the displaced 
nucleus of the corpuscle after the manner of a Halteridium ; 
it shows a very close resemblance to H. magna, found in the 
same hosts, but differs by the possession of a capsule. The 
parasite was named H. berestneffi by Castellani & Willey 
(1905), who, however, did not find the parasite in hosts of the 
same species from Ceylon. Dobell (1910) found the species 
in Hana tigrina from Ceylon. Patton (1908, p. 319) notes 
that he had the opportunity of studjang no less than five 
Hsemogregarines in E. tigrina and R. hexadactyla and also in 
the leech which transmits them, but he does not mention the 
species encountered. Wenyon (1926, p. 1398) records H. en- 
cajpsulse Berestneff as a parasite of R. limnocharis and both 
H. encajpsulse Berestneff and H. berestneffi Castellani & Willey 
from R. tigrina. The name H. encapsulse Berestneff is an 
error, as Berestneff did not give this name to his form, and 
so it must be treated as a synonym. 

Habitat. — Blood of Rana tigrina Daud. and R. limnocharis 
Wiegmann : Bombay, Bombay ; Rana tigrina Daud. : Ceylon, 
Colombo ; R. tigrina Daud. and R. hexadactyla Lesson, and 
in the leech which transmits them : Madras, Madras ; 
R. tigrina Daud. : PoRTUorESE India, Nova Goa ; also 
R. tigrina Daud. from India, in the Zoological Gardens, 



46. Haemogregarina cantliei Sambon & Seligmann. (Fig. 47.) 

■\Hxmogregarina sp., Laveran, 1905 (as noted in Wenvon, 1926, 

p. 1392). 
■\Hsemogregarina cantliei, Sambon & Seligmann, 1907 a, p. 1650 : 

1907 6, p. 310. 
■fHxmogregarina sp., Patten, 1908, p. 318. 

Hsemogregarina cantliei, Dobell, 1908, p. 293 ; Phisalix, 1913, 
pp. 304-7, 6 figs. ; Castellani & Chabners, 1919, p. 487 ; Wenvon, 
1926, p. 1392. 
Hsmogregarina sp., Wenyon, 1926, p. 1392. 

Adult sporonts club-shaped, the anterior third rounded or 
globular and the rest of body tapering to a slender tail ; en- 
closed within thick sausage-shaped capsules. Nucleus median 
or nearer posterior extremity, oval or irregular in outline and 
with chromatin frequently arranged in cross-strands. 

Fig. 47. — Heemogregarina cantliei Samb. & Selig. A, uninfected cor- 
puscle ; B, intra-corpuscular form surrounded by a capsule ; 
C, D, same without capsule ; E, free forms. (After Phisalix.) 

Dimensions. — Free sporonts, 16-18/x in length by 3-5jU, 
in breadth ; tail 2-3 /x in length ; nucleus 4/x long by 3/^ 

Habitat. — Blood of the snake, £'r?/a; corners Schneid. {=^Gongy- 
lophis conicus) : Madras, Madras. 

47. Hsemogregarina hankini Simond. (Fig. 48.) 

'\H8eniogregarina hankini, Simond, 1901 b, pp. 183-5 ; 1901 e, 

pp. 331-8, pi. vii. 
'fHeemogregarina sp. (? H. hankini Simond), Dobell, 1910, pp. 68, 79, 

pi. ii, figs. 9, 10. 
Hsemogregarina hankini, Castellani & Chalmers, 1919, p. 486 ; 

Wenyon, 1926, p. 1396. 

The parasite has two forms, vermicular and oval. The 
vermicular forms are large, doubled-up in the red blood- 
corpuscles in the circulating blood. Sometimes the two 
Hmbs approximately equal in thickness, sometimes one 
considerably thicker than the other. Nucleus a compact 
mass of deep, purple granules (Giemsa-stained). The oval 


forms also contain a nucleus which may be compact or frag- 
mented. Certain large forms are schizonts. 

Dimensions. — Vermicular form, length 12-15^ ; schizonts, 
20 ju, in diameter. 



Fig. 48. — Heemogregarina hankini Simond. A, large form with com- 
pact nucleus ; B, parasite with two nuclei, due to two 
parasites adhering together ; C, oval form, with a number 
of nuclei arranged on the periphery ; D, yoimg parasite 
with dispersed nucleus ; E, larger form with dispersed 
nucleus (n, nucleus of corpuscle ; p, parasite). (After 

Remarks. — According to Simond two forms of the parasite 
are met with, viz., vermicular and oval. In the young 
vermicular form the nucleus is compact or fragmented into 
a number of granules arranged in each limb of the parasite. 
The parasite is always doubled upon itself, the two limbs being 
parallel and sometimes in contact with one another. In the 
oval form also the nucleus may be compact or fragmented. 
Cells with incompletely developed merozoites were observed in 
the lung. These schizonts were spherical, and contained 
30 to 40 fusiform merozoites without any residue of segmenta- 
tion. If it be confirmed that schizogony takes place in the 
lung, the species will have to be transferred to the genus 

Habitat. — Blood of Gavialis gangeticus Gmelin : United 
Provinces, Jumna River ; Crocodilus porosus Schneider (?) : 
India (exact locality not cited) ; and Crocodilus porosus 
Schneider : Ceylon, Daduganoya, Veyangoda. 

48. Hsemogregarina laverani Simond. (Fig. 49.) 

■fHeemogregarina laverani, Simond, 1901 e, pp. 327-31, fig. 2. 
Hsemogregarina laverani, Castellani & Chalmers, 1919, p. 487 ; 

Wenyon, 1926, p. 1395. 
■\Hsemogregarina laverani, de Mello, 1932, pp. 1412-4, pi. i, 

figs. S 1-S 14. 

Young stages amoeboid, measuring barely 3/u. in diameter, 
provided with a small nucleus and many small cyanophil 
granules disposed either in the form of a coil or a horseshoe. 



Frequently the young stages are in the form of a minute 
vermicule bent upon itself, the Hmbs adhering in such a manner 
that it is difficult to distinguish this form from the amoeboid. 
Full-grown stage large, reniform, containing, besides the 
nucleus, two large refringent ovoid granules, and occupying 
a considerable extent of the surface of the corpuscle. The 
vermicular stage is much smaller than the reniform stage 
and does not extend beyond half the length of the corpuscle ; 
it possesses a bulging end and a short drawn-out tail, bent 
round on the swollen portion. 

Fig. 49. — Hsemogregarina laverani Simond. A, young amoeboid stage 
with a nucleus ; B, young stage in which the cyanophil 
grains are dispersed in the form of a coil ; C, young stage 
in which the cyanophil nuclear substance is in the form of 
a horseshoe ; D, young vermicular stage ; E, full-grown 
stage folded upon itself and showing two refringent granules. 
(From de Mello, after Simond.) 

Remarks. — The chief characteristic of the species is the 
possession in most of the stages of two refringent granules, 
which serve to distinguish this species from all others. 

Habitat. — Blood of Lissemys punctata granosa (Schoepff) : 
United Provinces, Agra. 

49. Haemogregarinaleschenaultii Robertson. (Fig. 50.) 

■\Hsemogr6garina leschenaultii, Robertson, 1908, pp. 182, 184, figs. 4-7. 
Hsemogregarina leschenaultii, Wenyon, 1926, pp. 1388. 

Hsemogregarine with two free motile forms, only present 
in the blood : (1) slender free form with dense nucleus ; 
no granules in the cytoplasm ; rather actively motile : 
(2) broad, massive, granular form ; less active, period of move- 
ment succeeded by period of rest. Two intra-corpuscular 
forms also present : (1) long recurved form corresponding 
exactly with the slender free form ; causes hypertrophy of 
the corpuscle ; this is always the prevailing type in an in- 
fection : (2) broad form with reticulate nucleus, growing 
to a somewhat larger size ; never very numerous even in a good 



Dimensions. — Free motile forms, length 26-28 /x ; broad, 
intra-corpuscular form, 30 /x. 

Habitat.— 'Eloo^ of the lizard, Hemidactylus leschenaulti 
Dum. & Bibr. : Ceylon, Trincomalee. 


Fig. 50.^ — Hsemogregarina leschenaultii Robertson. A, broad intra- 
corpuscular form ; B, broad free form ; C, slender recurved 
intra-corpuscular form ; D, slender free form. (After 

50. Hsemogregarina magna (Grassi & Feletti). (Fig. 51.) 

Hasmogregarina ranarum (part), Kruse, 1890, p. 541 ; Celli & 

Sanfelice, 1891, p. 504, pi. v, figs. 16-18. 
Drepanidium magnum, Grassi & Feletti, 1891, p. 82 ; 1892, pi. i, 

fig. 15. 
Drepanidium krusei, Labbe, 1892, p. 617. 
Danilewskya krusei, Labbe, 1894, p. 127. 
Hsemogregarina magna, Labbe, 1899, p. 76. 

■\Danilewskya krusei, Berestneff, 1903, p. 347, pi. viii, figs. 9-13. 
Hsemogregarina magna, Patton, 1908, p. 319 (?) ; Wenyon, 1926, 

p. 1398. 

Adult form oval, very large, may be folded upon itself in 
the corpuscle. 

Habitat. — Blood of Rana tigrina Daud., R. limmocharis 
Wiegmann : Bombay, Bombay ; R. tigrina Daud. : Madras, 

51. Hsemogregarina malabariea de Mello. (Fig. 52.) 

"f Hsemogregarina inalaharica, de Mello, 1932, pp. 1411-25, pi. ii, 
figs. A-D. 

Two forms are present : (1) The vermicular form consists 



of a sausage-shaped body with a caudal appendage which is 
always apphed to the body along its concave border. The 
convex border is bounded by a thin band of protoplasm which 




Fig. 51. — Hsemogregarina magna (Grassi & Feletti). A, young intra- 
corpuscular form ; B, adult oval form ; G, adult form folded 
upon itself; D, uninfected corpuscle. Uninfected corpuscles 
are also sliown with A, B, and G for the sake of comparison. 
(After Berestneff. 

Fig. 52. — Hsemogregarina malabarica de Mello. A, free vermicular 
form ; B, intra-corpuscular vermicular form ; G, intra- 
eorpuscular bean-shaped form ; D, cyst, containing eight 
merozoites ; E, F, larger merozoites found in the liver cell. 
A, B, G as seen in the blood ; D, as seen in smears from the 
liver; E,F, as seen in sections of the liver. (After de Mello.) 

stains a clear blue or sometimes violet with Romanowsky's 
stain, and is thicker at the pole at which the nucleus is situated, 
becoming narrower to form the tail ; the tail does not extend 


to the other pole of the body. The nucleus is nearly always 
found near the zone of separation of the two parts, and 
contains irregularly dispersed chromatin particles. The body 
is more or less filled with metachromatic granules. The 
parasitized corpuscles become hypertrophied, measuring 20- 
25 fi, whereas the normal corpuscles measure 16-1 8 ju,. (2) The 
reniform or bean-shaped forms are found in very much smaller 
numbers than the vermicular ones. The nucleus is rounded 
and central, but may sometimes be situated close to one of the 
poles ; it contains chromatin granules or rods. The cyto- 
plasmic constitution is also different from that of the vermicular 
form. In one of the poles there are two or more vacuoles ; 
there may be vacuoles, but fewer, in the other pole also. The 
parasitized corpuscle is hypertrophied, but is enlarged in its 
width rather than in its length, measuring 17-18;u, by 14-16/x. 
Schizogony takes place in the hver, and is of two types, 
resulting in large and small merozoites. Cysts are found 
in the liver of the host ; they are rounded, with a thick 
membrane, and contain 8 rounded or falciform merozoites. 
The free merozoites are generally found in pairs. In sections 
of the Hver the cysts containing 8 merozoites are not en- 
countered, but larger merozoites, 12-16/x in size, occur singly, 
or in groups in 2, 4, 8 or larger numbers. 

Dimensions. — Vermicular form 17-22 fju, reniform type 
8-12 ju, ; cysts, diameter 18-20/^ ; merozoites 6-8 fx or 12-16|U,. 

Remarks. — This species is recognizable from H. laverani 
by the young stages not being amoeboid and the vacuoles 
being inconstant in number and occurring at one or both 
poles of the reniform stages ; the vermicular stage also differs 
in being larger than the reniform stages, in occupying nearly 
the whole of the corpuscle, and in having a tail nearly as long 
as the body of the parasite. The species resembles H. vittatss 
in possessing cysts with 8 merozoites, but in H. malabarica 
the cysts are regularly spherical, whereas in H. vittatee they 
are navicular and the reniform stages of that species contain 
so-called plastids. Schizogony in the lungs or in the blood 
does not occur in this species, though it is known to take place 
in others. 

The species also somewhat resembles H. nicorise, but is 
clearly marked off from it by the non-existence of a clear pole, 
the chromatic granules filling completely the opposite pole, 
the tail being closely applied to the body, the vermicular 
forms being larger and more abundant than the bean-shaped 
forms, and by the chromatoid granules filling up the cyto- 

Habitat. — Blood, liver, bone-marrow and other viscera of 
the Indian tortoise, Lissemys punctata granosa (Schoepff) : 
Portuguese India, Nova Goa. 



52, HsBmogregarina mesnili Simond. (Fig. 53.) 

1901a, pp. 150-2; 1901 e, 
1919, p. 487; 

"fHcemogregarina mesnili, Simond, 
pp. 322-7, fig. 1. 
Hsemogregarina mesnili, Castellani & Chalmers, 
Wenyon, 1926, p. 1395, 

The young stages are amoeboid. The older stages may be 
either vermicular with a distinct horn -like prolongation bent 
upon the main body, oval or reniform, slender and doubled, 
or folded into three parts. The nucleus may be fragmented 
or compact. 

Dimensions. — In the folded forms the total length of the 
parasite may exceed 30 /a. 


Fig. 53, — Hsemogregarina mesnili Simond, A, young amoeboid stage ; 
B, horned vermicular stage, showing two nuclei ; G, adult 
reniform stage with colourless granules ; D, vermicular 
stage, with two equal limbs ; E, very long vermicular stage, 
folded twice over, with three limbs interlacing together ; 
F, fragmentation of a schizont, with free merozoites. (After 

Remarks. — The young amoeboid forms measure 3-6 ^a in 
length and contain a fragmented nucleus. The horned vermi- 
cule stage recalls the appearance of Hsemoproteus metchnikowi, 
described by the same author from Chitra indica. The reni- 
form individuals are often seen to be filled with refringent 
colourless granules. The elongate vermicular forms are 
uniform in diameter throughout the greater part of their length, 
and are folded into two equal limbs, or in the largest individuals 
folded twice over, the three limbs interlacing like a figure of 8 , 
a characteristic not met with in other species. 

Habitat.— Blood of Kachuga tectum (Gray) : United 
Pkovincbs, Jumna River. 

53. Hsemogregarina mirabilis Castellani & Willey, (Fig. 54.) 

^Hsemogregarina mirabilis, Castellani & Willey, 1904, pp. 86-90,. 

figs. 28-38 ; Patton, 1908, p. 318. 
Hsemogregarina mirabilis, Dobell, 1908, p. 294, 
■^Hsemogregarina jnirabilis, Plimmer, 1913, p. 149. 
Hsemogregarina mirabilis, Castellani & Chalmers, 1919, p, 487, 
fig. 153 ; Wenyon, 1926, p. 1394. 
•\Hsemogregarina mirabilis, Scott, 1926, p. 236. 

Form elongate, thick, Gregarine-like and bent ; cytoplasm 


(with Romanowsky's stain) stains uniform blue, leaving no 
clear pole ; the nucleus is stained reddish-blue and is near 
the anterior pole. 

Fig. 54. — Hsemogregarina inirabilis Cast. & Will. A, intra-corpuscular 
form bent double ; B, parasite within the capsule inside 
a corpuscle ; G, D, parasite emerging from the cyst and the 
corpuscle ; E, F, free forms. (After Gastellani & Willey . ) 

Dimensions. — Length about 12 ju,. 

BemarJcs. — Gastellani and Willey (1904) described the 
occurrence within the corpuscles of parasites of relatively 
large size and shghtly crescentic or reniform shape, and con- 
sisting of a membrane {cytocysl) inside which developed an 
elongate body {monozoite) with a well-defined nucleus. 
This organism escaped from the cytocyst and the corpuscle 
and became a freely motile organism in the blood. The 
exact stages cannot be determined from their description, 
but probably they were dealing with the development of the 
gametes from the gametocytes. 

Habitat. — Blood of Tropidonotus asperrimus Bouleng. : 
€eylon ; and Tropidonotus piscator Schneider : Ceylon (also 
in specimens from Ceylon in the Zoological Gardens, London) : 
Madras, Madras. 

54. Hsemogregarina najse Laveran. (Fig. 55.) 

Hsemogregarina najse, Laveran, 1902, p. 1037, figs. 1-3. 
^Hsemogregarina najse, Patton, 1908, p. 318. 
Hsemogregarina najse, Dobell, 1908, p. 293. 
■\Hxmogregarina najse, Plinuner, 1912, p. 413 ; 1913, p. 149 ; 1914, 
p. 188 ; 1916, p. 85. 
Hsemogregarina najse, Gastellani & Chalmers, 1919, p. 487; Wenyon, 

1926, p. 1110, pi. xix, figs. 8-10 ; p. 1393. 
■fHsemogregarina najse, Scott, 1926, p. 236 ; 1927, p. 189. 

Intra-corpuscular form elongate, vermicular, rounded at 
one end and drawn out at the other. When fully developed, 
bent upon itself. Nucleus oval, more or less elongate, 
situated about the middle. 

Dimensions. — ^Vermicular form 14ju, in length ; fully 
developed form 21-22ju. by 3 /x at the rounded end. 



Remarks. — -Schizonts of this species were discovered by 
Wenyon (1909) in the lungs of Naja hajx in the Sudan. 

Fig. 55. 

-Haemogregarina najaa Laveran. A, elongated form ; B, re- 
curved form ; G, form liberated from the corpuscle. (After 

Habitat. — Blood of Naja naja (Linn.) : Madras, Madras 
(also in specimens from India in the Zoological Gardens, 

55. Haemogregarina nicorise Castellani & Willey. (Fig. 56.) 

■fHsemogregarina nicorise, Castellani & Willey, 1904, pp. 85-6, figs. 17- 
27 ; Patton, 1908, p. 319 ; Robertson, 1908, p. 178 ; 1910^ 
pp. 741-62, pis. xxxii-xli ; Dobell, 1910, p. 68 ; Alexeieff, 
1912, p. 97. 
Hsemogregarina nicoriee, Castellani & Chalmers, 1919, p. 486 ;: 
Wenyon, 1926, pp. 1084, 1105, 1395 ; Reichenow, 1929, p. 927. 

Form elongate, Gregarine-like, with one end granular and 
the other end clear, the central nucleus being a more or less 
diffuse aggregation of chromatin granules. Sometimes the 
organism is bent round upon itself. Schizogony takes place 
in the blood-vessels of the lung and produces nearly 70 mero- 
zoites, or in the circulating red blood- corpuscles producing six 
to eight smaller merozoites which grow into gametocytes. 

When the Hsemogregarines are taken together with the 
blood of the tortoise into the crop of the leech, some of them 
pass into the intestine and are there found as motile vermicules. 
They penetrate into the intestinal wall, where the differentia- 
tion of the hitherto indistinguishable gametes takes place, 
culminating in a process suggesting anisogamous conjugation* 
The zygote breaks up to form eight sporozoites, which pass 
through the intestinal wall into the blood-spaces. The 
Hsemogregarine is probably passed into the blood of the tortoise 
through contamination of the wound by the leech while 

Dimensions. — Length 12^. 

Remarks. — The Hfe-history of the organism, as worked out 
by Robertson, confirms in all essential respects that described 
by Reichenow (1910) for H. stepanowi. In H. nicorias 




schizogony, producing the large merozoites, occurs free in the 
blood-vessels of the lung, and that producing the smaller 
gametocytes in the circulating red blood-corpuscles. In 
H. stepanowi, on the other hand, schizogony always takes 
place in the bone-marrow, always inside the blood-corpuscles, 
and the number of merozoites does not exceed twenty-four. 

Christophers once showed Patton some bodies from a leech 
off Lissemys punctata granosa which suggested developmental 
forms of the Hsemogregarine of the tortoise, but Patton came 
to the conclusion that they probably represented some stage 
in the hfe-cycle of a Coccidium parasitic in the leech. 

Fig. 56. — Hsemogregarina nicorias Castellani & Willey. A, bean-shaped 
Hsemogregarine, with circular type of nucleus ; B, early 
stage of schizogony in the blood-stream ; G, late stage of 
schizogony in the lung ; D, fully-formed merozoites, only 
a very few of the total number formed shown in the section ; 

E, free motile stage in the lumen of the intestine of the leech ; 

F, microgametocyte lying closely applied to macrogameto- 
cyte ; G, the microgametocyte giving rise to the micro- 
gamete nuclei, one of which fuses with the nucleus of the 
macrogamete ; H, sporocyst with eight nuclei; /, sporocyst 
showing sporozoites. (After Robertson.) 

Habitat. — Blood of the tortoise Geoemyda trijuga (Schweiger) 
and in the leech Ozobranchus shipleyi Harding : Ceylon, 
Colombo ; blood of the tortoise Lissemys punctata granosa 
(Schoepfif) and in the transmitting leech : Madras, Madras. 



56. Hsemogregarina nueleobisecans Shortt. (Fig. 57.) 

■fHcBmogregarina nueleobisecans, Shortt, 1917, pp. 408-12, pi. xxxi, 

figs. 1-17. 
Hasmogregarina bisecans, Wenyon, 1926, p. 1397. 
■\H3em0gregarina sp., Donovan (firstrecorciedin Wenyon, 1926, p.l397). 
■fHiemogregarina sp., Wenyon, 1926, p. 1397 ; Scott, 1926, p. 236. 

Fig. 57. 

G H 

-Hsemogregarina nueleobisecans Shortt. 


schizont; B, division of schizont; 
merozoites ; D, E, single infection 

F, double infection of an erythrocyte : , 

an erythrocyte; H, parasite passing out of ruptured 
erythrocjdie ; /, parasite free in plasma. (After Shortt.) 


A, mature imdivided 
(7, cyst, containing 
of an erythrocyte ; 
G, treble infection of 


Schizogony only in the liver. The mature schizont broadly 
oval and lying in a thin- walled cyst closely surrounded by 
hepatic cells. The cyst grows larger, large number of nuclei 
are produced, and a correspondingly large number (some- 
times as many as 115) of very small merozoites are produced. 
Each merozoite has a pink-staining, hyaline protoplasm 
and a well-marked nucleus. Gametocytes intra -corpuscular, 
sausage-shaped, slightly curved, and lying with concave 
border applied to the nucleus of the host- cell. Sometimes 
a gametocyte may He with its convex border towards the 
nucleus, or may occupy one end of the corpuscle. Sometimes 
one or both extremities are effilated and slightly recurved. 
The gametocytes may vary considerably in size : each is 
enclosed in a thick capsule ; the cytoplasm stains (with 
Leishman's stain) an azure-blue or faint pink colour, and 
contains a large ovoid nucleus, staining a dull crimson. Two 
or three gametocytes may be found in the same corpuscle. 
Free forms are found in the plasma, having escaped by 
rupture of the corpuscle, leaving the empty capsule behind. 
Sporogony not known. The secondary host may perhaps 
be a blood-sucking worm, Angiostoma sp., found in the lung 
of the toad, or a sand-fly. 

Dimensions. — Mature schiznot 16/i, by 7/x ; fully developed 
cyst 28-30 ju, by 24-5-26 ju, ; merozoite 3/x by l-3ju, ; gameto- 
cytes from 9-5 ju. by 4 /x to 21-8 ju, by 4-8 /i. 

Habitat. — Blood of Bufo melanosticus Schneider : Punjab, 
Ambala : Delhi ; United Provinces, Cawnpore ; also from 
the same host from India in the Zoological Gardens, London. 

57. Hsemogregarina pythonis (Billet). 

Danilewshya pythonis. Billet, 1895, p. 30, figs. 1-3. 

H^mogregarina pythonis, Labbe, 1899, p. 76. 
■fHsemogregarina pococki, Sambon, 1907, p. 283. 
^Hsemogregarina sp., Patton, 1908, p. 318. 

Heeinogregarina pococJci, Dobell, 1908, p. 293. 

Hsemogregarina pythonis, Johnstone, 1912, p. 23.5. 
JHse^nogregarina pococki, Plimmer, 1912, p. 412 ; 1913 p 148 - 

1914, p. 189; 1916, p. 85; 1917, p. 32. 
jHrnmogregarina sp., Phisalix, 1913, pp. 1052-4. 

Hiemogregarina pythonis, Castellani & Chalmers, 1919, p. 487. 

Hsemogregarina sp., Wenyon, 1926, p. 1393. 
■fHeemogregarina pococki, Wenyon, 1926, p. 1393 ; Scott, 1926, p. 236. 

Intra-corpuscular. Body club-shaped, often folded in the red 
blood- corpuscle. Anterior extremity broader and rounded. 
Posterior extremitj^ attenuated and recurved. Cytoplasm 
more or less granular. Nucleus median or nearer posterior 
extremity, large, oval, and with coarse, deeply staining 
chromatin granules. Parasite lies parallel or obliquely to 
long axis of host-cell, of which it occupies about two-thirds. 



without causing much alteration beyond displacement of 

Dimensions. — 14-16 /u. in length. 

Remarks. — Plimmer found the host-ceUs often deformed and 
generally diminished in size. Johnstone (1912) regards 
H. pococki Sambon as identical and synonymous with 
H. pythonis (BUlet). 

Habitat. — Blood of Python molurus (Linn.) : Madras, 
Madras ; also in specimens from India in the Zoological 
Gardens, London. 

58. Hsemogregarina rara Laveran & Mesnil. (Fig. 58.) 

fHsemogregarina rara, Laveran & Mesnil, 1902, p. 611. 
Haemogregarina rara, Castellani& Chalmers. 1919, p. 487 ; Wenyon, 
1926, p. 1395. 

Intra-corpuscular or free. The intra -corpuscular form is 
elongate or curved into the shape of an arc, with one of the 
extremities rounded and the other more or less attenuate. 
Sometimes the parasite lies in one part of the corpuscle without 
displacing its nucleus, at other times it is strongly recurved, 
and the nucleus of the corpuscle is pushed to one end. When 
the organism is liberated from the corpuscle it is fusiform, 

A B 

Fig. 58. — Hsemogregarina rara Laveran & Mesnil. A, elongated ; 
B, curved form ; C, form leaving the corpuscle ; D, E, free 
in the plasma. (After Laveran and Mesnil.) 

transparent, and motile, and possesses a very elongate and 
clear nucleus. In the stained preparation (with Leishman's 
stain) the cytoplasm is of a light blue colour and is finely 
granular, with large chromatoid bodies. The nucleus is 
elongate, cyhndrical, nearly always swollen at the ends, 
and is stained deep violet. It occupies about two-thirds of 
the length of the body. Multiplication appears to be by binary 
fission. Other stages of development have not as yet been 

Dimensions. — Average length 15 ju by 2-3 /x in width. 

Habitat. — Blood of the tortoise, Ghinemys reevesii (Gray) : 



59. Hsemogregarina rodriguesi de Mello, de Sa, de Sousa, 
Dias, & Noronha. (Fig. 59.) 

-\Hgemogregarina rodriguesi, de Mello, de Sa, de Sousa, Dias, & 
Noroiiha, 1917, pp. 13, 14, pi. i, figs. 8-14; de Mello, 1934 a, 
p. 1786. 

rHmmogregarina proencee, de Mello, de Sa, de Sousa, Dias, & Noronha, 
1917, pp. 14, 15, pi. i, figs. 15-17 ; de Mello, 1934 a, p. 1786. 

The parasite occurs in two forms, elongate or shorter and 
stumpy. In the extra-corpuscular form it is gregariniform 
and without a capsule. Cytostome stains blue with Leishman's 
stain, and the nucleus is elongate, rectangular, generally 
compact, rarely vesicular, occupying a large part of the 
parasite and situated near one or other extremity. The 
intra -corpuscular stage has the same appearance, but possesses 
a large clear capsule containing sometimes one or two vacuoles 
in the centre. The short and stumpy form is equally broad 
at both poles, and has a less extensive capsule. 





Fig. 59. 

-Hsemogregarina rodriguesi de Mello, de Sa, de Sousa, Dias, 
& Noronha. A—D, intra-corpuscular forms ; E, free form. 
(After de Mello and others.) 

Remarks. — The parasite is very common, 50 per cent, of 
the lizards examined being infected. It could not be deter- 
mined whether schizogony takes place in the blood or in the 
internal organs. The short stumpy form was originally 
described as a separate species under the name of H. proencse, 
but de Mello (1934 a) has amalgamated the two. 

Habitat. — Blood of Hemidactylus hrooki Gray : Poettjguese 
India, Nova Goa. 

60. Hsemogregarina stepanowiana Laveran & Mesnil. (Fig. 60.) 

f Hsemogregarina stej)anowiana, Laveran & Mesnil, 1902, pp. 609—10, 
figs. 1-5. 
Hsemogregarina stepanowiana, Castellani & Chalmers, 1919, p. 487 ; 
Wenyon, 1926, p. 1395. 

Intra-corpuscular form oval or reniform, displacing the 
nucleus when fully developed ; cytoplasm finely granular and 
containing numerous chromatoid particles ; nucleus central 



rounded or oval, placed at right angles to the long axis of the 
organism, and containing chromatin granules of various 
sizes. When the parasite doubles upon itself the two portions 
are not equal, and the nucleus is always situated in the longer 
part near the level of the bend. Extra-corpuscular form like 
a vermicule, with one extremity much wider than the other. 
The intra-corpuscular form, when about to multiply, assumes 
an oval form, and its nucleus divides repeatedly into eight 
nuclei ; the protoplasm then divides into as many parts, 

Fig. 60. — Hsemogregarina stepanowiana Laveran & Mesnil. A, B, 
elongated corpuscular forms ; C, fuU-gi'own parasite doubled 
upon itself ; D, free vermicule-like form ; E, form showing 
nuclear multiplication. (After Laveran and Mesnil.) 

the corpuscle breaking up at the same time. Multiplication 
forms were not found in the blood, but were numerous in 
smears from the liver. 

Dimensions. — Extra-corpuscular form 18-20|U, by S/x. 

Remarks. — According to the authors of the species the young 
form shows greatest resemblance to the young form of 
H. stepanowi Danilewsky ; but the mature form differs markedly 
from that species. In H. stepanowi the parasite becomes 
doubled exactly about its middle, its nucleus is very elongate, 
and the nucleus of the corpuscle is displaced, but not hyper- 
trophied. The Kberated Hsemogregarine is a vermicule, 
measuring 30-40 ju, by 3-4 /z. In H. stepanowiana the doubling 
of the intra-corpuscular parasite is not exact, and the extra- 
corpuscular vermicule is much shorter and wider than in 
H. stepanowi. 

Habitat. — -Blood of the tortoise, Chinemys reevesii (Gray) 

61. Hsemogregarina testudinis Laveran & Nattan-Larrier. 

'\Hsemogregarina testudinis, La^^eran & Nattan-Larrier, 1912, 
pp. 134^6, 10 figs. (Abstract in Wiegmann's Archiv, 1913, p. 411). 
Hsemocystidiurn testudinis, Castellani & Chalmers, 1919, p. 516. 
Hsemogregarina testudinis, Wenyon, 1926, p. 1396. 

Small, medium, and large intra-corpuscular forms. Free 


and multipKcation forms also present. Cysts 12-15 /i in length 
and 12-14 ft in width. Number of merozoites in each cyst 
4 to 8 macro- or 12 to 24 microgametes. 

Habitat. — Blood of the tortoise, Testudo emys Schleg. & Miill. : 
India (exact locahty not cited). 

€2. Haemogregarina thomsoni Minchin. (Fig. 61.) 

^Heemogregarina thomsoni, Minchin, 1907, pp. 1098-1104, pis. Iv-lvi, 

figs. 1-23. 
Heetnogregarina sp., Shortt, 1917, pp. 402-8, pi. xxx, figs. 1-21. 
Heemogregqrina thomsoni, Castellani & Chalmers, 1919, p. 487 ; 

Wenyon, 1926, p. 1388. 

Both intra-corpuscular and free forms (so-called vermicules) 
are met with. Intra-corpuscular parasite always distinctly 
sausage-shaped, and slightly bowed in the plane of the corpuscle, 
this normal ciu-vature being due to the parasite being situated to 
the side of the corpuscular nucleus. In a few cases the curve 
of the parasite is reversed, its convexity being towards the 
corpuscular nucleus, which is then much more displaced, and 
the parasite resembles its free form very closely. 

Young intra-corpuscular forms vsiTy in length from about 
a half to two-thirds of the blood- corpuscle ; cytoplasm 
very clear, and a dehcate nucleus, consisting of faintly 
staining granules and strands of chromatin, forms a band 
round the waist of the parasite. Full-grown intra-cor- 
puscular parasites are at least three -fourths the length 
of the blood-corpuscle ; cytoplasm finely granular, nucleus 
exceedingly rich in chromatin, forming a deeply staining mass 
of irregularly spongy texture, occupying the middle region of 
the body for its whole width and nearly one-third of its length. 
The free vermicules closely resemble the full-grown intra- 
corpuscular forms, long, medium, and stumpy forms being 
distinguishable . 

Sometimes most of the intra-corpuscular parasites have the 
form of an elongated sausage, slender and drawn out ; cyto- 
plasm clear and free from coarse granulations ; nucleus forming 
a band or zone at the middle of the body, equal in width to 
nearly half the length of the body, and with chromatin arranged 
in the form of transverse strands. 

Small forms are generally situated at one end of the erythro- 
cyte ; larger forms He in the long axis of the host-ceU, and displace 
the nucleus laterally. The parasite is usually situated with its 
concave border embracing the convexity of the erythrocyte 
nucleus, but its position may be reversed. Invasion by two 
or more forms is not uncommon, the parasites lying parallel 
to one another and both on the same side, or the two embracing 



the nucleus between them. Infected cells, especially those 
with two parasites, are increased in size. 

Schizogony takes place in the endotheUal cells of the 
capillaries of the lung, and is of two forms. One form of 
schizont produces a smaller cyst, with larger and less numerous 
merozoites, and the other form produces a larger cyst, with 
smaller and more numerous merozoites. In the former case 
the schizont in the endotheHal cell is broadly oval, and usually 
forms a smaller cyst giving rise to 16 merozoites. The larger 
cysts are more rare, and may contain as many as 40 merozoites. 
What the two kinds of merozoites develop into has not been 
determined. The process probably differs from that described 

Fig. 61. — Hsemogregarina thomsoni Minchin. A, small iiitra-corpusciilar 
form ; B, large intra-corpuscular form showing the so-called 
" capsule " ; C, a large and an intermediate-sized form iia the 
same corpuscle ; D, two large individuals lying side by side, 
both on one side of the nucleus ; E, two large individuals 
embracing the nucleus between them ; F, the free form. 
(After Minchin.) 

by Reichenow and Robertson for the Haemogregarines of 
cold-blooded Vertebrates. 

Dimensions. — Intra-corpuscular forms, young 9-11 /x in 
length, full-grown 15-17 ft by 5 ju. ; schizonts 15 /m by 6 /x ; 
cysts, smaU 21-5 )u, by 14-5 fi, large 25 /x by 15 /j,. 

Habitat. — ^Blood of the lizard, Agama tuberculata Gray : 
Punjab, Kasauh. 



63. HaBmogregarina thyrsoidese de Mello & Vales. (Fig. 62.) 

'\IIgemogregarina thyrsoidese, de Mello & Vales, 1936, pp. 403-4, 

pi. XXXV. 

Youngest form ovoid, with conspicuous nucleus and cyto- 
plasm stained light blue with Romanowsky's stain. The full- 
grown parasite shows irregular vacuoHzation of the cytoplasm. 
The nucleus is generally central, often divided into two more 
or less irregular masses, with indistinct nuclear membrane, 



Fig. 62. — Hsemogregarina thyrsoidem de Mello & Vales. A, young form ; 
B, normal full-grown form ; G, form with central nucleus, 
containing two chromatin granules ; D, form with one 
nuclear mass at each pole. (After de Mello and Vales.) 

and minute volutin granules. Sometimes the chromatin 
is reduced to two very minute centrally situated granules ; 
or the nucleus is situated at one of the poles and composed 
of two masses, or the two nuclear masses are situated one at 
each pole. 

Habitat. — Blood of the freshwater eel, Thyrsoidea macrurus 
(Bleeker) : Portuguese India, Nova Goa. 

64. Haemogregarina triedri Robertson. (Fig. 63.) 

■[Hsemogregarina triedri, Robertson, 1908, pp. 181, 184, figs. 10, 11. 
Hsemogregarina iiedri, Wenyon, 1926, p. 1388. 

Fig. 63. 

-Hsemogregarina triedri Robertson. A, broader bean-shaped 
form ; B, long form. (After Robertson.) 



Haemogregarine with double capsule, a delicate inner capsule 
and a loose outer one, with tendency to stain very deeply with 
Giemsa's stain. The inner capsule has an opercular lid at 
one end. Two forms present : (1) broad, rather bean-shaped 
form ; (2) long, slightly recurved specimens, with elongate 
nuclei. Both forms frequently show at one end an 
irregularly shaped body which stains a bright red with 
Giemsa's stam. The young forms have no capsule. 
Transmitting host not known. 

Dimensions. — 13-1 5 |U, in length. 

Habitat. — Blood of the lizard, Hemidactylus triedrus 
(Daudin) : Ceylon, Trincomalee. 

65. Haemogregarina vittatse Robertson. (Fig. 64.) 

"fHeemogregarina vittatee, Robertson, 1908, pp. 179, 180, 183, figs. 2-3. 
Hsemogregarina vittatse, Castellani& Chalmers, 1919, p. 487, figs. 149, 

157, 158. 
'\Hssmogregarina vittatee, Donovan (first recorded in Wenyon, 

1926, p. 1395). 
■\Hg&mogregarina vittatse, de MeUo, 1932, pp. 1417-18, pi. i, figs. Rl, 


Hsemogregarine infection associated almost mvariably with 
Trypanosoma vittat^. Two forms present : (1) broad massive 
form ; (2) recurved form mth pale cytoplasm, the two limbs 

Fig. 64. 

-Hsemogregarina vittatse Robertson. A, elongate, recurved 
form ; B, broad massive form. (After de Mello.) 

being equally long ; nucleus dense. Broad form shows reticu 
late dense cytoplasm and rather delicate loose nucleus ; the 
larger forms have two red-staining plastid (?)-lLke bodies at 
one end. Schizogony occurs in the spleen and Hver ; eight 
merozoites are formed ; these are enclosed in pairs in a 
delicate boat-shaped capsule. Transmitting host probably 

Dimensions. — Recurved form 22-26 /x in total length. 

Habitat. — Blood of the tortoise, Lissemys punctata granosa 
(SchoepiBF.) : Ceylon ; Portuguese India, Nova Goa. 


66. Haemogregarina xavieri de Mello. (Fig. 65.) 

■\Heemogregarina xavieri, de Mello, 1932, pp. 1426-27, pis. iii & iv. 

Two forms present : (1) Broadly ova] or sometimes reniform, 
occupying, according to the state of its development, a greater 
or less extent of the corpuscle. Cytoplasm alveolar, containing 
generally two, sometimes more, rounded vacuoles, usually 
confined to the polar regions. Nucleus central, containing 
granules or rods of chromatin in compact masses or variously 
dispersed. The parasitized corpuscles are generally not 
altered, sometimes reduced or enlarged in size, and the nucleus 
of the corpuscle is displaced to one side. (2) Elongate or 
vermicular forms, sometimes enclosed in a fine, membranous 
capsule or entirely without a capsule, provided with a tail 
which stands out from the body, and does not exceed one- 
quarter or one-third of its length. These tailed forms are 
much less numerous than the oval forms. One pole sometimes 
presents one or two vacuoles, but generally the c3rtoplasm 
alveolar, and the vacuoles not clearly indicated. Nucleus 
central, and the chromatin showing the same disposition as in 
the oval forms. The metachromatic granules generally 
abundantly distributed in the tail, and rarely in the anterior 
pole as well. Forms intermediate between the oval and the 
vermicular are also met with, and sometimes two individuals 
are seen in the same corpuscle. 

Schizogony takes place in the lungs, spleen, liver, etc. 
Cysts found in smears from the lungs are oval, and contain 
2 or 3 schizonts ; those found in smears from the spleen are 
rounded or oval, and contain 6 or 7 schizonts ; while those 
in smears from the hver are circular or more or less oval, 
and contain 2 or 3 schizonts. In sections of parasitized organs 
oval cysts are found to contain 6 or 7 schizonts (merozoites) 
similar to those met with in the smears. Fusiform merozoites 
are seen in the blood-stream and in the intercellular spaces. 

Dimensions. — Oval form 7-5-13-5/a in length by 2-5-6-2/x 
in width ; vermicular form 9-10//, in length by 4ju. in width ; 
cysts from lungs 8-10 ju, by 4-5 ju,, from hver, spherical, 15 ju, in 
diameter, or oval, 8-5//, by 5/x or lOjU, by 7-5 jU. 

Remarks. — The species is morphologically distinct from 
H. laverani Simond, described from the same host at Agra. 
Both H. malabarica and H. xavieri do not show any amoeboid 
young stages, nor do they show the two refringent granules 
so characteristic of H. laverani. H. xavieri differs from 
H. malabarica in the tail being considerably shorter, and not 
being closely appHed to the body, the nucleus being central in 
the elongate forms, and in the dimensions being considerably 
smaller in both elongate and oval forms than in the other 


Fie 65— H^mogreqarina xavieri deWello. A In blood ^4, i?, o^a 
'g- '°^- \^^^. ^c. kidney-shaped form ; D, elongate form with 
a short tail : E, elongate form withm a capusle. 

Si cyst containLg few schizonts ; B, cyst contammg 
six sehi;onts ; C, cyst six schizonts (^erozo^tes) ; 
D E fusiform merozoites m the blood ; A, B, 0, m smears 
frim lungs, spleen, and liver respectively ; D, E, m section 
of a blood-vessel . ( After de Mello . ) 


Habitat. — Blood, lungs, spleen, and liver of the tortoise, 
Lissemys punctata granosa (Schoepflf) : Pobtuguesb India, 
Nova Goa. 

In addition to the species that are recorded above, a number 
of Hsemogregarines have been observed in a variety of hosts ; 
but, unfortunately, the published descriptions are so meagre 
that it is impossible to refer the forms to any known species. 
For the sake of completeness I have given these records below. 

67. Haemogregarina sp. 

'fHsemogregarina sp., Plimmer, 1914, p. 189. 
Heemogregarina sp., Wenyon, 1926, p. 1403. 

Habitat. — Blood of the fish, Trichogaster fasciatus Schneider : 
from India, in the Zoological Gardens, London. 

68. Haemogregarina sp. 

■\Hse7n0gregarina sp., Simond, 1901 e, p. 320; Plimmer, 1912, p. 412; 

1914, p. 189. 
Hs&tnogregarina sp., Wenyon, 1926, p. 1390. 

Large ; of ordinary type. Nucleus of parasitized cell divided 
by the parasite into two parts with a connecting thread. 
Schizogony in lungs. 

Remarks. — This form may be the same as H. varani Laveran, 
1905, described from Varanus niloticus. 

Habitat. — Blood of the lizard, Varanus monitor (Linn.) : 
India ; also from India, in the Zoological Gardens, London. 

69. Haemogregarina sp. 

Hsemogregarina sp., Dobell, 1908, p. 292. 
■fHiemogregarina sp., Patton, 1908, p. 318 ; Plimmer, 1913, p. 149. 
Haemogregarina sp., Wenyon, 1926, p. 1391. 

Both short and long forms present. 

Habitat. -S\oodL oi Bungarus cseruleus (Schneid.) ( =Eungarus 
candidus) : Madras, Madras ; also from India, in the Zoo- 
logical Gardens, London. 

70. Haemogregarina sp. 

^Hsemogregarina sp., Robertson, 1908, p. 182, fig. 12. 

Hsemogregarina sp., Dobell, 1908, p. 294. 
1[H^mogregarina sp., Patton, 1909, pp. 149, 152 ; Dobell, 1910, p, 70 ; 
Plimmer, 1912, p. 413 ; 1913, p. 149 ; 1914, p. 188. 

Haemogregarina sp., Wenyon, 1926, pp. 1391, 1394. 

Young intra-corpuscular form without a capsule. Full-grown 
form, with a very marked capsule thickened at both ends, and 
showing a deep red staining area at either end and a delicate 
nucleus. Free motile form also without a capsule. 


Remarks. — Five species are known from various species 
of Zamenis from other parts of the world, and the forms 
described by Robertson and Dobell and noted by Patton and 
Plimmer may belong to one of them. 

Habitat. — Blood of the snakes, Chrysopelea ornata (Shaw) : 
Ceylon ; and Zaocys mucosus (Linn.) : Ceylon, Colombo, 
Paradeniya ; Madras, Madras ; also in same host from India, 
in the Zoological Gardens, London ; in the Imguatihd, Poro- 
cephalus pattoni Stephens, from the lung of Zaocys mucosus 
(Linn.) : Madras, Madras. 

7L Haemogregarina sp. 

'^Heemogregarina sp., Plimmer, 1914, p. 189. 
Hsemogregarina sp., Wenyon, 1926, p. 1391. 

Habitat. — Blood of the snake. Coluber helena Daud., from 
Ceylon, in the Zoological Gardens, London. 

72. Haemogregarina sp. 

■\Hsemogregarine, Simond, 1901 e, p. 320. 
Heemogregarina sp., Dobell, 1908, p. 293 ; Wenyon, 1926, p. 1391. 

Habitat. — Blood of a snake, Coluber sp. : India. 

73. Haemogregarina sp. 

^Heemogregarina sp., Dobell, 1910, p. 70. 

Habitat. — Blood of the snake, Dipsadomorphus forstenii 
{Dumeril & Bibron) : Ceylon, Colombo. 

74. Haemogregarina sp. 

'\Hsemogregarina sp., Dobell, 1910, p. 70. 

Habitat. — Blood of the snake, Dipsadomorphus ceylonensis 
Gunther : Ceylon, Paradeniya. 

75. Haemogregarina sp. 

■\ Haemogregarina sp., Patton, 1908, p. 318. 
Haemogregarina sp., Dobell, 1908, p. 293 ; Wenyon, 1926, p. 1391. 

Habitat. — Blood of the snake, Dendrophis pictus GmeUn : 
Madras, Madras. 

76. Haemogregarina sp. 

■fHsem,ogregarina sp., Patton, 1908, p. 318. 

Hsem,ogregarina sp., Dobell, 1908, p. 293. 
^Hsemogregarina sp., Dobell, 1910, p. 70. 

Hsemogregarina sp., Wenyon, 1926, p. 1392. 

Habitat. — Blood of the snake, Dryophis mycterizans (Daud.) : 
Madras, Madras ; Ceylon, Colombo, Paradeniya. 


77. HsBmogregarina sp. 

^Hsemogregarina sp., Patton, 1908, p. 318. 

Hseniogregarina sp., Dobell, 1908, p. 293. 
fHsemogregarina sp., Plimmer, 1912, p. 414 ; 1913, p. 149 ; Scott. 
1926, p. 236. 

Hsemogregarina sp., Wenyon, 1926, p. 1392. 

Habitat. — Blood of the snake, Eryx johnii (Russ.) : Madras, 
Madras ; also from India, in the Zoological Gardens, London. 

78. Hsemogregarina sp. 

■\Heemogregarina sp., Plimmer, 1913, p. 149 ; Scott, 1926, p. 236. 
Haemogregarina sp., Wenyon, 1926, p. 1393. 

Habitat. — Blood of the snake, Naja bungarus Schleg., from. 
India, in the Zoological Gardens, London. 

79. Hsemogregarina sp. 

jHxmogregarina sp., Simond, 1901 e, p. 320 ; Robertson, 1908, 
p. 182. 

Hsemogregarina sp., Dobell, 1908, p. 293. 
^Hsetnogregarina sp., Dobell, 1910, p. 70. 

Hsemogregarina sp., Wenyon, 1926, p. 1393. 

Remarks. — ^The form probably belongs to H. najse. Laveran, 
described from the same host. 

Habitat. — Blood of the snake, Naja naja var. atra (Cantor) : 
India ; blood of Naja naja (Linn.) : Ceylon, Paradeniya. 

80. Hsemogregarina sp. 

^Hsemogregarina sp., Simond, 1901 e, p. 320. 
Hsemogregarina sp., Wenyon, 1926, p. 1393. 

Habitat. — Blood of a snake, Naja sp. : India. 

81. Hsemogregarina sp. 

^Hsemogregarina sp., Robertson, 1908, pp. 182, 183. 
Hsemogregarine sp., Wenyon, 1926, p. 1393. 

Hsemogregarine showed extraordiaary activity. The free 
form moved with a rapid swimming motion, and entered 
a blood- corpuscle by simply piercing it, swimming round 
between the nucleus and the corpuscle wall, and bursting the 
corpuscle by curling and then suddenly straightening itself. 
The process takes a few seconds. It also injures the corpuscles, 
which it touches in passing, the corpuscle losing all its haemo- 
globin immediately. The intra-corpuscular stage very closely 
resembles the free form. 

Remarks. — Possibly the form examined by Robertson may 
have been the same as H. pythonis (Billet). 

Habitat. — Blood of a snake, Python sp. : Ceylon. 


82. Haemogregarina sp. 

Laverania sp., Billet, 1895, p. 31. 

Hsemogregarina sp., Labbe, 1899, p. 77. 
'fH£emogrega7-ina sp., Patton, 1908, p. 318. 

" Danilewskya '' =Hx?nogregarina sp., Dobell, 1908, p. 294. 
fHsemogregarina sp., Dobell, 1910, p. 70. 

Hsemogregarina sp., Wenyon, 1926, p. 1394. 

Remarks. — Dobell (1910) thinks that it is probably the same 
as " Danilewskya " described in this host from Tong-king by 
Billet (1895) {==H£emogregarina sp., Labbe). 

Habitat. — Blood of Tropidonotus stolatus (Linn.) : Madras, 
Madras ; Ceylon, Paradeniya. 

83. Hsemogregarina sp. 

■^Hsemogregarina sp., Patton, 1908, p. 318. 

Hseinagregarina sp., Dobell, 1908, p. 294. 
■\Hsemogregarina sp., Plimmer, 1912, p. 414 ; 1913, p. 149 ; 1915, 
p. 130. 

Hsemogregarina sp., Wenyon, 1926, p. 1394. 

Habitat. — Blood of Russell's Viper, Vipera russellii (Shaw) : 
Madras, Madras ; also from India in. the Zoological Gardens, 

Genus HEPATOZOON Miller, 1908. 

Leucocytozoon, James, 1905 a, pp. 1-12 ; 1905 b, p. 1361 ; Bentley, 

1905, p. 988. 
Hsemogregarina, Balfour, 1905 a, p. 240 ; 1905 b, p. 1330. 
Hsemogregarine, Laveran, 1905, p. 295. 
Leucocytozoa, Patton, 1906, pp. 1-13 ; 1908, p. 319. 
Leucocytozoon, .). R. Adie, 1906, p. 325 ; Christophers, 1906, 

pp. 1-16 ; 1907, pp. 1-12. 
Hepatozoon, Miller, 1908, pp. 1-48, 
Leucocytozoon, Porter, 1908, pp. 703-16. 
Leucocytogregarina, Porter, 1909, p. 264 

Hsemogregarina, Wenyon, 1910, pp. 70-1 ; 1911, pp. 324-32. 
Leucocytozoon, Christophers, 1912, pp. 37-44. 
Hepatozoon, Minchin, 1912, pp. 372, 376. 
Leucocytogregarina, Sangiorgi, 1912, pp. 287-93 ; de Mello, de Sa, 

de Sousa, Dias, & Noronha, 1917, p. 13. 
Hepatozoon,Wenyon, 1926, pp. 1085-6 ; Reichenow, 1929, pp. 920-3 ; 

Kudo, 1931, p. 281 ; Calkins, 1933, pp. 361, 527. 

Schizogony takes place in cells of the iaternal organs (bone- 
marrow, liver, spleen, kidney) of Vertebrates. After several 
generations of merozoites have been produced some of them 
enter erythrocytes or leucocytes and develop into gametocytes. 
These are taken into the body of a blood-sucking Arthropod 
(tick, mite, louse, etc.) , and the micro- and macrogametes develop 
and unite ia pairs. The zygote becomes encysted in the oocyst, 
which increases enormously in size, eventually producing 
sporoblasts, sporocysts, and sporozoites. Each large oocyst 




contains numerous sporocysts, all of which produce numerous 

The genus was founded by Miller for a parasite of the leuco- 
cytes of rats, which had been previously referred to the genera 
Hsemogregarina or Leucocytozoon. 

84. Hepatozoon adiei Hoare. (Fig. 66.) 

"fHepatozoon adiei, Hoare, 1924, pp. 63-6, 1 pi. 
Hepatozoon adiei, Wenyon, 1926, pp. 1086, 1095, fig. 456 ; Reiche- 
now, 1929, p. 923. 

Schizonts more or less regularly ovoid. Nuclear multiplica- 
tion gives rise to 12 to 24 nuclei. Merozoites elongate and 
vermiform. Younger schizonts enclosed in definite cells, 



IFig. 66. — Hepatozoon adiei Hoare. A, B, ovoid schizonts ; C, schizont 
showing formation of merozoites ; D, rod-like gametocyte 
enclosed in a leucocyte. A-C, in the smears of the lung ; 
D, in the peripheral blood. (After Hoare.) 

probably endothelial cells of the capillaries. In later stages, 
host-cell reduced to a thin membrane with the flattened 
nucleus at one side of it. Gametocytes rod-Hke, rounded 
at both extremities ; nucleus usually terminal in position. 
The gametocyte either displaces or is wedged in the leucocyte 

Dimensions. — Schizonts from 8/x by 4-8 /u- to 20-8 /x by 11-2)li ; 
gametocytes 8-5 /x by 4-5 /x. 

Remarks. — Only dried films of the blood and smears of the 
internal organs of the bird were available for study. The 
schizonts were found in smears of the lung, and the gametocytes 
in the leucocytes of the peripheral blood. The schizonts and 


the gametocytes closely resemble H. canis and H. muris. 
Nothing is known of the sexual cycle in the Invertebrate. 

Habitat. — Blood and smears from the lung of an Indian 
eagle (not identified) : Punjab, Kasauli. 

85, Hepatozoon canis (James). (Fig. 67.) 

■\Leucocytozoon canis, James, 1905, pp. 1-12, 1 pi. ; Bentley, 1905, 
p. 988; Christophers, 1906, pp. 1-16, 1 pi.; 1907, pp. 1-12, 1 pi. 
'fLeucocytozoon bentleyi, James, 1905 a, p. 1361. 
Leucocytogregarina rotundata, Patton, 1910, p. 280. 
Heemogregarina canis adusti, Nuttal, 1910, p. 108. 
Hasmogregarina canis, Wenyon, 1911, pp. 324-32, text-figs. 4-8, 

pi. xvi, figs. 1—17. 
Leucocytozoon canis, Christophers, 1912, pp. 37-4:4:, 1 diagram, 

text-figs. 4-8. 
Hepatozoon canis, Minchin, 1912, p. 377. 
Heemogregarina canis, Castellani & Chalmers, 1919, p. 483. 
^Hepatozoon canis, Rau, 1925, p. 293. 
\Hsemogregarina canis, Rau, 1926, p. 244. 
Hepatozoon canis, Wenyon. 1926, pp. 1091, 1092, fig. 455 ; pi. xix, 
figs. 3, 4 ; pp. 1355-7 ; Reichenow, 1929, pp. 921, 922, figs. 885, 

Schizogony occurs in the spleen, bone-marrow or liver. 
Schizonts of several types ; some produce a small number 
(generally three) of large merozoites, which become schizonts 
again, whilst others produce a large number of small merozoites. 
These latter enter the mononuclear or polymorphonuclear 
leucocytes and become gametocytes. Free vermicules found 
in the gut of the infected ticks. 

The gametocytes undergo development in the common 
dog-tick, Ehipicephalus sanguineus Koch, in the tissues of 
which are eventually produced very large oocysts, containing 
from 30 to 50 sporocysts, each containing about 16 sporozoites 
and a residual body. After the sporocysts are fully developed, 
the oocyst appears to break up and sporozoites are found 
scattered among the tissues of the tick. Infection of the dog is 
probably produced by the ticks being eaten. 

Dimensions. — Gametocytes 8-10 ^u, by 4-5 /x, nucleus 2-3 ft; 
oocysts up to 100 ju, ; sporocysts 15-16 /a in length ; sporozoites 
14-15 /A in length. 

Remarks. — Rau (1925) infected dogs by injection of spleen 
material containing schizonts, as also by the injection of the 
tissues of infected ticks. Parasites appeared in the blood 
two or three weeks after inoculation and infection progressed 
in intensity, bringing about the death of some of the animals. 
Wenyon, however, doubts if other causes of death were 

Habitat. — Internal organs and leucocytes of the Indian wild 
dog, Gyon duhhunensis (Sykes) : Assam, BorjuHe, Gauhati, 




Fig. 67. — Hepatozoon canis (James). A, parasite in a leucocyte of the 
dog ; B, free vermicule in the blood of the dog ; C, D, 
schizogony in the bone-marrow of the dog ; E, gametocytes 
lying side by side ; F, G, formation of the microgametes ; 
H, zygote showing a number of nuclei ; /, section of oocyst 
in which sporoblast formation is taking place ; J, section of 
fully developed oocyst in which the sporocysts, each con- 
taining about sixteen sporozoites, have formed. A-G in the 
dog ; H-J in the tissues of the tick. (A-I, after Christophers ; 
J, after Wenyon.) 


Tejpur, Nowgong ; the dog, Canis familiar is (Shaw) : Madras, 
Madras ; the jackal, Canis aurem Linn. : Madras, Madras ; 
the fox, Vulpes bengalensis (Shaw) : Madras, Madras. Also 
the tick, Bhipicephalits saTiguineus Koch : Madras, Madras. 

86. Hepatozoon felis domestici (Patton). 

'\Leitcocytozoon felis domestici, Patton, 1908, p. 319. 
Heemogregarina felis, Castellani & Chalmers, 1919, p. 486. 
Hepatozoon felis, Wenyon, 1926, pp. 1085, 1355. 

Remarhs. — Patton (1908) merely recorded this as a new 
species, without giving any description. It is, therefore, not 
certain that this is morphologically distinct from H. canis. 

Habitat. — Leucocj^es of the cat, Felis sp. : India. 

87. Hepatozoon funambuli (Patton). (Fig. 68.) 

jLeucocytozoon funambidi, Patton, 1906, pp. 1—13, pi. i ; 1908, p. 319. 
Hepatozoon funambuli, Minchin, 1912, p. 377. 
Heemogregarina funambuli, Castellani & Chalmers, 1919, p. 486. 
Hepatozoon funamhuli, Wenyon, 1926, pp. 1085, 1358. 

The gametocyte has an elongate oval body, with one end 
larger and rounder than the other. It lies in the large mono- 
nuclear leucocyte, and shows slow vermicular movements. 
The narrow end shows a distinct bend upwards, simulating 
a tail. In escaping from the host-cell the more rounded 
end protrudes first. The free vermicules are elongated and 
spindle-shaped ; they are very active, performing rapid 
serpentine movements, twisting and curling about. In 
preparations stained with Romanowsky's stain cytoplasm 
stains light blue, somewhat darker at the two ends. There is 
no capsule round the parasite, but by prolonged staining a faint 
pink outline is seen round it. Nucleus large, irregularly 
quadrilateral, lying about the centre, staining more deeply 
but not uniformly. A number of large chromatic dots are 
seen in the cytoplasm, more in one extremity than in the other. 
Sometimes two parasites are seen in the same leucocyte, 
lying close to each other, between the two detached parts of 
the nucleus, or one on either side of the nucleus. The free 
vermicule contains a rounded or oval nucleus. 

Dimensions. — Intracorpuscular forms 10 /x by 5jU. in the 
widest part ; free vermicules 13-14 /x by 3-4 /x. 

Remarks. — The parasites were present in the peripheral 
blood and in smears from the spleen and Hver. Vermicules 
were also seen from the mid-gut and the body- cavity of the 
louse, Hsematopinus, but no developmental forms were met 



Habitat. — Leucocytes of the squirrel, Funambulus pennantii 
Wroughton, and the body of the louse, Hasmatopinus sp, : 
Bombay, Kathiawar. 

Fig. 68. — Hepatozoon funambuU (Patton). A, two leucocytes, showing 
two parasites in each ; B, three free forms, showing chromatic 
particles ; C, four free parasites from spleen-smears ; D, free 
attenuate form from peripheral blood ; E, free vermicule 
from the peripheral blood ; F, vermicule from the mid-gut of 
the squirrel-louse ; G, large vermicule from the body-cavity 
of the louse. (After Patton.) 

88. Hepatozoon gerbilli (Christophers). (Fig. 69.) 

^Hsemogregarina gerbilli, Christophers, 1905, pp. 1-15, 1 pi. 
Hsemogregarina gerbilli, Patton, 1906, p. 3. 
Hepatozoon gerbilli, Minehin, 1912, pp. 376, 377. 
Hsemogregarina gerbilli, Castellani & Chalmers, 1919, p. 485. 
Hepatozoon gerbilli, Wenyon, 1926, pp. 1085, 1358 ; Reichenow, 
1929, p. 922. 

Stages of the schizogonous multiplication are not known. 
Gametocytes are found in the red blood-corpuscles and are 
marked by a short hook-shaped projection at their hinder end. 



Nucleus close to the bent end. Active vermicule stage free 
in the plasma. Further development takes place in the louse, 
Heemafopinus stephensi. In the ahmentary canal of the louse 
free worm-like parasites are recognized. In the body- cavity 
are found large oocysts in various stages of development. 
The fully developed oocysts contain numerous oval sporocysts ; 
each sporocyst contains 6 to 8 sporozoites and a residual body. 

Fig. 69. — Hepatozoon gerbilli (Christophers). A, forms in the red 
blood-corpuscles ; B, fully matured oocyst ; C, sporocyst con- 
taining sporozoites. (After Christophers.) 

Dimensions. — Oocysts 350 /x in diameter. 

Remarks. — In imperfectly stained films the general appear- 
ance of the parasite is very like that of the malarial crescents, 
except that the pigment is absent. This is due to the " tail " 
not being differentiated from the body. 

Habitat. — ^Red blood- corpuscles of the Indian gerbil, 
Tatera indica (Hardw.) {^Gerbillus indicus), and in the mid-gut 
and the pyloric ampulla of the louse, Hsematopinus stephensi 
Christophers & Newstead : India (locahty not cited). 



Fig. 70. — Life-cycle of Hepatozoon muris (Balfovir). A, sporozoites 
escaping from sporocysts in the intestine of rat ; B, sporozoites 
penetrating through intestinal epithelium and entering 
into the blood-vessels of the villi ; C, sporozoites passing 
from blood-vessels into liver-cells ; D-G, schizogony in liver- 
cells ; H, merozoites escaping from liver-cells and entering 
other liver-cells to repeat schizogony; I, merozoites (gameto- 
cytes) leaving liver-cells to enter blood-vessels ; ./, gameto- 
cytes in mononuclear cells of blood; K, gametocytes 


89. Hepatozoon leporis (Patton). 

■fLeucocytozoon leporis, Patton, 1908, p. 319. 
Hepatozoon leporis, Wenyon, 1926, p. 1359. 

Patton (1908) merely recorded this as a new species, without 
giving any description. 

Habitat. — Blood of the black-naped hare, Lepus nigricollis 
(Cuvier) : Madras, Madras. 

90. Hepatozoon muris (Balfour). (Fig. 70.) 

Leucocytozoon muris, Balfour, 1905, pp. 110-11, pi. xi. 
^Leucocytozoon ratti, Adie, 1906, pp. 325-6, text-figs. 

Hepatozoon perniciosiim. Miller, 1908, pp. 1-48, pis. 

Hepatozoon muris, Minchin, 1912, pp. 376, 377. 

Heemogregarine ratti, Castellani & Chalmers, 1919, p. 486. 

Hepatozoon tnurts, Wenyon, 1926, pp. 1085, 1086-90, fig. 453, pi. xix, 

figs. 1, 2. 
'\Hepatozoon muris, Donovan (recorded in Wenyon, 1926, p. 1360). 

Hepatozoon muris, Reichenow, 1929, pp. 920-1 ; Kudo, 1931, 
p. 282, fig. 119, A;. 

Schizogony takes place in the hver-cells of a rat. Smallest 
schizont is a spherical, uninucleate body. The schizont 
increases in size, and the nuclei multiply by repeated division 
till there are 12 to 20. The full-grown schizont is surrounded 
by a delicate cyst-wall. Merozoites are budded off, enter 
other cells, and repeat schizogony. After a time the merozoites 
become young gametocytes, enter the blood-vessels, invade 
the mononuclear leucocytes, and appear as Hsemogregarines. 
When the blood is sucked by the mite, Leelaps echidninus, 
the Hsemogregarines are hberated from the leucocytes and 
escape from the enclosing cysts. The gametocytes associate 
in pairs, each becoming flattened to produce an elongated 
body, with pointed extremities. Complete fusion of the two 
gametocytes is said to take place. Before actual union takes 
place the macrogametocyte increases somewhat in size and 
encloses the smaller microgametocyte. After fertilization 
the zygote elongates and becomes a motile ookinete ; this 
moves about in the stomach-contents, increasing in size. 
Later it penetrates through the intestinal wall and settles 
down in the surrounding tissue, becoming spherical and growing. 

escaping from mononuclear cells in the stomach of the mite ; 
L-0, syngamy and penetration of intestinal wall by zygote 
(ookinete) ; P—S, growth of zygote (sporont) in oocyst 
in tissues of mite ; T, surface of sporont, showing sporoblast 
formation by budding ; U, portion of oocyst containing 
sporoblasts ; V, portion of oocyst containing sporoblasts 
with multiplying nuclei ; W, portion of oocyst containing 
sporocysts in each of which are a number of sporozoites. 
The mite is eaten by the rat, in the intestine of which the 
sporozoites escape from the sporocysts. (From Wenyon, 
after Miller.) 


It acquires a cyst-wall and grows still further to become 
the ripe oocyst. Sporogony takes place in the body-cavity 
of the mite. Nuclear multipUcation and formation of sporo- 
blasts and sporocysts take place, each spore containing 
12 to 24 sporozoites. Re-infection takes place by a rat 
devouring infected mites. 

Dimensions. — Schizont, young 10 /it, full-grown 30-35 ju, by 
25-28 ft ; ookinete, young 25 /x by 10 /a, full-grown 50 /* by 25 /a ; 
oocyst 200-250^. 

Remarks. — According to Miller (1908) and the Japanese 
workers Kusama, Kasai, and Kobayashi (1919), complete 
fusion of the two gametocytes takes place, and no formation 
of micro- or macrogametes is described. Wenyon (1926), 
however, from what is known of other Hsemogregarines, 
such as H. stepanowi, thinks it probable that the micro- 
gametocyte produces microgametes, and that a fertilization 
of the Adelea type occurs instead of the complete fusion of the 
two gametocytes. 

Miller succeeded in infecting rats by placing on them 
infected mites, and also by contaminating food with crushed 
mites, and as injection of crushed infected mites into the peri-^ 
toneal cavity did not produce infection, it is obvious that 
infection takes place by way of the alimentary canal. 

Habitat. — Leucocytes of the brown rat, Rattus norvegicus 
(Berkenhout) {=^Mus decumanus=Mus norvegicus) : Punjab ; 
Rattus rufescens (Gray) {=Mus rufescens) ; and the rat-mite, 
Leelaps echidninus Berlese : India. 

91. Hepatozoon sp. 

•\Hepatozoon sp., Donovan (recorded in Wenyon, 1926, p. 1362). 
Habitat. — Blood of the flying squirrel, Pteromys petaurista 
Pallas : India. ' 

Genus KARYOLYSUS Labbe, 1894. 

Heemogregarina (part), Danilewsky, 1886. 

Karyolysus, Labbe, 1894, p. 109 ; 1899, p. 75 ; Minehin, 1903, 

pp. 261, 265. 266 ; 1912, p. 372 ; Reichenow, 1913, pp. 317-63 ; 

1921, pp. 179-291 ; Wenyon, 1926, pp. 1095-101 ; Reichenow, 

1929, pp. 865, 916-19 ; de Mello & de Meyrelles, 1937, pp. 119- 


Sporogony in the epitheHal cells of the intestine of the 
Invertebrate host, a mite, produces an oocyst with a number 
of sporoblasts, which escape from the oocyst as motile 
vermicules (sporokinetes) and enter the egg of the mite, where 
they secrete sporocysts within which sporozoites are developed. 
The mite hatched from the egg has the sporocysts in its 
intestinal epithehum. The sporocysts enter the intestinal 



epithelium of the Vertebrate host, a lizard, through the 
ingestion of the faeces of the nymph or of the nymph itself. 
The sporozoites pass to the blood-vessels and penetrate the 
endotheUal cells, where schizogony takes place. Certain 
merozoites enter the red blood- corpuscles as gametocytes 
and appear as Hsemogregariaes. The gametocytes are taken 
up by the mite, in the gut of which gametogony takes place 
and the oocyst is formed. 

Bemarks. — The genus owes its names to the fact that the 
nucleus of the host-cell is often karyolysed and fragmented, 
but this character cannot be regarded as of generic value. 

92. Karyolysus jorgei de Mello & de Meyrelles. (Fig. 71.) 

■\Karyolysus jorgei, de Mello & de MeyreUes, 1937, pp. 119-41, pi. vi, 

4 text-figs. 

Gametocytes, occurring in the red blood- corpuscles in the 
peripheral circulation of the lizard, are elongate oval, marked 

Fig. 71. — ^Life-cycle of Karyolysus jorgei de Mello & de Meyrelles. 

A, male and female gametocytes in peripheral blood ; 

B, endoglobular cycle in capillaries producing macro- 
cysts with four merozoites ; G, endothelial cycle in liver 
and lungs producing microcysts with eight leishmaniform 
merozoites ; D, free merozoites which may invade either . 
the red cells or endothelial cells. (After de Mello and de 

by a strong limititig membrane, and contain a large central 
nucleus, a smaller roundish paranuclear body, and near the 
opposite pole a more or less irregular zone named as a polar 
capsule. The gametocytes are sexually differentiated. In 


the male the nucleus is central, oval or quadrangular in form, 
and always compact. In the female the nucleus may often 
be polar, and always contains a definite karyosome in its centre. 
In the capillary circulation occurs the endoglobular cycle. 
A merozoite invades the red corpuscle, grows into a schizont, 
and by twice repeated binary division forms a macrocyst 
containing four large gregariniform merozoites. These mero- 
zoites invade the endothelial cells of the lungs and hver and 
produce microcysts, each containing eight leishmaniform 
merozoites. The microcysts hberate the merozoites either in 
the interstices of the tissues or in the protoplasm of the endo- 
thehal cell itself. This second schizogonic cycle has been 
designated as the endothelial cycle. The leishmaniform. 
merozoites may invade either the red corpuscles, where they 
undergo binary divisions again, or the endothehal cells, where 
they again become microcysts. It is not known from what 
particular forms the gametocysts are derived. Sporogony 
completely unknown. 

Dimensions. — Gametocytes measure 5-11 fx by 1-5-2-5 [x. 
Macrocysts measure 13-15 /x by 8-1 Oju,; macromerozoites 
measure 8-1 l/i. by 2-5-3jLt. Leishmaniform merozoites: 
roundish 1-5-3 ju, in diameter, fusiform 3-5^ by 1-1-5 /x. 

Remarks. — The lack of karyolitic action on the part of the 
parasite and the occm-rence of endoglobular schizogony makes 
the organism a remarkable transitional form between H8em,o- 
gregarina and the previously known species of Karyolysus. 
The differentiation of the gametocytes and the occurrence of 
a paranuclear body and specially differentiated polar area in 
the gametocytes and the pecuhar form and structure of the 
micromerozoites are other characteristic features of the 

Habitat. — Red blood- corpuscles and endothelial cells of the 
liver and lungs of Calotes versicolor Daud. subsp. major Bl3^h : 
Portuguese India, Nova Goa. 

II. Suborder EIMERIDEA Leger, 1911. 

The schizonts develop into micro- and macrogametocytes 
which are similar in size and develop independently of one 
another. The microgametocyte produces a relatively large 
number (six or more) of microgametes. The motionless zygote 
secretes a resistent oocyst, which does not increase in size. 
The asexual and the sexual cycles occur in the same host. 

Various schemes of classification of the suborder have been 
proposed. It is generally admitted that the composition of the 
mature oocyst, viz., the number of sporocysts and sporozoites 


it contains, provides the most convenient diagnostic characters 
for the differentiation of the genera : but different opinions 
have been held regarding the basis upon which the genera 
should be united into subfamiHes and famihes. Following 
Ltihe (1906), Minchin (1912), Reichenow (1921) and Wenyon 
(1926) arrange the genera into higher groups, mamly according 
to the type of schizogony. Schneider (1881) and Leger (1911) 
adopted the characters of the ripe oocyst as the basis for classi- 
fication : but while Schneider and his followers, Biitschli 
(1882), Labbe (1899) and Minchin (1903), arranged them 
according to the number of sporocysts within the oocyst, 
Leger (1900), foUowed by Mesnil (1903), Poche (1913), Doflein 
(1916), Pinto (1928), and Noller (1928), classified them according 
to the total number of sporozoites in the oocyst. 

Wenyon (1926) divides the suborder into six famihes, viz., 
Selenococcidiidse, Cryptosporidiidse, Eimeriidse, Caryotrophidse, 
Aggregatidae, and Lankesterelhdae. Reichenow (1929) re- 
shuffled the genera among three famihes, and brought over 
Dobelhdse from the Adeleidea and Leucocytozoidse from the 
H^MOSPOKiDiA. Thus he classified the Eimeridea into five 
famihes, viz., Selenococcidiidge, Aggregatidse, Dobellidse, 
Eimerudse, and Leucocytozoidae. 

Hoare (1933), leaving out of consideration the famihes 
Dobelhdse and Leucocytozoidse (which might as well be 
included, as in Wenyon's classification), has proposed a modi- 
fication of Leger's classification, in which the subfamilies 
are distinguished from each other by the number of sporocysts 
within the oocyst, while the genera within each subfamily 
differ from one another in the number of sporozoites within 
each sporocyst. The suborder is divided into two famihes : 
(1) Selenococcidiidse and (2) Eimeriidse. The family Eimeriidse 
is divided into six subfamilies. I have followed this classifica- 
tion in this work. 

Identification Table of Families. 

1 (2). Body cylindrical or vermiform. Nuclear 

multiplication takes place in the extra- 
cellular motile stage ; schizont becomes 
rounded on entering an epithelial cell 
and breaks up into eight merozoites, 
which are set free into the limien of the 
gut. Numerous microgametes formed 

from a microgametocyte. Fertiliza- [Leger & Duboscq. 

and sporogony unknown Selenococcidiidse * 

2 (1). Body not cylindrical or vermiform Eimeriidse Leger, em. 

[Hoare, p. 158. 


1. Family EIMERIID^ Leger, 
emend. Hoare, 1933. 

Schizogony and sporogony are very uniform in character 
throughout the family, but variations occur in the number of 
the sporocysts and the sporozoites developed within the 
oocyst. The family is divided into six subfamilies on this 
latter basis. 

Key to Subfamilies and Genera. 

I. Schizogony and sporogony in the same host. 

1. Oocyst asporocystid Subfam. Cryptosporidiin^ 

[Poche, emend. Hoare, p. 159. 

Oocyst tetrazoic Cryptosporidium * Tyzzer. 

Oocyst octozoic Pfeijferinella * Wasielewski. 

Oocyst octozoic Schellackia * Reichenow. 

Oocyst polyzoic Lankesterella Labbe, p. 159. 

2. Oocyst monosporocystid Subfam. Cabyosporin^ * 


Sporocyst tetrazoic Mantonella * Vincent. 

Sporocyst octozoic Caryospora * Leger. 

3. Oocyst disporocystid Subfam. Cyclosporins 

[Wenyon, p. 162. 

Sporocysts dizoic Cyclospora * A. Schneider. 

Sporocysts tetrazoic Isospora A. Schneider, p. 162. 

Sporocysts octozoic Dorisiella * Ray. 

4. Oocyst tetrasporocystid Subfam. Eimeriins Wenyon. 

[p. 173. 

Sporocysts dizoic Eimeria A. Schneider, p. 173. 

Sporocysts tetrazoic TFewi/oneZZa Hoare, p. 197. 

Sporocysts polyzoic Angeiocystis * Brasil. 

5. Oocysts octosporocystid Subfam. Yaeimovellins * 

Sporocysts polycystid Yakimovella * Gouseff. 

6. Oocyst polysporocystid Subfam. Barroussins 

[(Wenyon), p. 199. 

Sporocysts monozoic Barroussia * A. Schneider. 

Sporocysts monozoic Echinospora * Leger. 

Sporocysts tetrazoic Pythonella Ray & Das- 

[Gupta, p. 199. 

II. Schizogony and sporogony in different hosts. 

Oocysts polysporocystid Subfam.AGGREGATiNsReiche- 

[now, emend. Hoare, p. 199. 

Sporocysts dizoic Merocystis * Dakin. 

Sporocysts dizoic Pseudoklossia * Leger & 


Sporocysts trizoic Aggregata Frenzel, p. 200. 

Sporocysts dodecazoie Caryotropha * Siedlecki. 

Sporocysts dodecazoie Ovivora * Mackinnon & Ray. 

Sporocysts polyzoic Myriospora * Lermantoff. 


Subfamily CRYPTOSPORIDIIN^ Poche, 
emend. Hoare, 1933. 

Oocyst contains four, eight, or many sporozoites developed 
without the formation of sporocysts. 

Genus LANKESTERELLA Labbe, 1899. 

Drepanidium, Lankester, 1882, pp. 53-65 ; Labbe, 1894, p. 76. 

Lankesterella, Labbe, 1899, pp. 73-5 ; Minchin, 1912, pp. 372, 378 ; 
N6]ler,19136,p. 231; 1920 6, p. 176; Wenyon, 1926, pp. 878-80, 
1105; Eoiowles, 1928, p.471; Reichenow, 1929, pp. 964-5; Kudo, 
1931, p. 275 ; Calkins, 1933, p. 566 ; Hoare, 1933, pp. 383, 385. 

The whole of the development takes place in the endothelial 
cells of the blood-vessels. The oocyst contains numerous 
(32 or more) sporozoites developed without the formation 
-of sporocysts. The sporozoites finally enter the blood- 
corpuscles and are mechanically transferred to another host 
by a blood-sucking animal. 

Remarks. — This genus includes certain types which were 
originally considered as Haemogregarines of cold-blooded 
animals. Noller (1913 a, 6 and 1920 h) showed that the forms 
within the blood-corpuscles are sporozoites, and that the rest 
of the cycle takes place in the endothehal cells of the blood- 
vessels (instead of the intestine as in the typical coccidian) 
and is of the Eimeria type. Reichenow (1919) described 
another genus, Schellackia, from Hzards, in which the develop- 
ment is on similar Hues, but takes place in the intestine. 
He (1921 a) placed the two genera in a new family, Lankesterel- 
lidse, and Wenyon (1926) followed this arrangement ; but 
Reichenow (1929) has since placed both genera in the family 
Eimeriidse, and Hoare (1933) has put them in a new subfamily, 
-Cryptosporidiinse, of the same. 

Key to Indian Species. 

Sporozoite like a vermicule, with its anterior [p. 159. 

extremity tapering, 10-15 fi in length ... L. minima (Chaussat), 
Sporozoite usually constricted into 3 segments, 

the middle containing the nucleus; [p. 162. 

15-16 /x in length L. monilis (Labbe), 

53. Lankesterella minima (Chaussat). (Fig. 72.) 

Anguillula minima, Chaussat, 1850. 

Drepanidium ranarum, Lankester, 1871, pp. 387-9, figs. 3, 4; 

1882, pp. 53-65; Labbe, 1891, p. 479; 1892, p. 617: 1893', 

p. 1207. 


Drepanidium princeps, Labbe, 1894, p. 76. 

Lankesterella ranarum, Labbe, 1899, p. 74 ; Minchin, 1903, pp. 239, 

253-7, 260, 261, 265, 267, 270, 345 ; 1912, pp. 372-8. 
Lankesterella minima Minchin 1903 p. 265. 
'\ Lankesterella minima, Patton, 1908, p. 319 ; 1909, pp. 146-7. 
Hsemogregarina minima Mathis & Leger, 1911, pp. 448-9. 
Lankesterella minima, Minchin, 1912, p. 372 ; Noller, 1912, pp. 201-8, 

pi. xx; 1913 a, pp. 313-16; 1913 6, pp. 222-32, pi. xiv, figs. 55- 

68 ; pi. XV, figs. 69-72 ; 1920 a, pp. 169-89, pis. iv-vi; Wenyon, 

1926, pp. 878-80, fig. 380 ; p. 1105. 
■\ Lankesterella minima, Scott, 1926, pp. 237-8 ; 1927, pp. 190-1. 
Lankesterella minima, Reichenow, 1929, pp. 964-5, fig. 930 ; Kudo, 

1931, p. 276, fig. 15, g. 
Lankesterella ranarum. Calkins, 1933, p. 545, fig. 218, B, C. 

Infection of the frog is brought about by a leech, which 
introduces the sporozoites. Sporozoites make their way into 
the blood- capillaries of various organs, and apparently enter 
the endothehal cells, where the entire development takes place. 
Each sporozoite becomes rounded, grows into a schizont, 
and produces a large number of merozoites. The merozoites 
escape into the blood and infect other endothehal cells. Mero- 
zoites of a special kind are finally produced, and these, after 
entering the endothehal cells, develop into micro- and macro - 
gametocytes. Microgametocyte produces a large number of 
microgametes, and fertilization of the macrogamete results. 
An oocyst is formed round the zygote, which breaks up directly, 
without the formation of sporoblasts and sporocysts, into 
a number of sporozoites. The latter, by rupture of the oocyst, 
escape into the blood and enter red blood-corpuscles. Here 
the sporozoite is seen as a small vermicule, which may attain 
half the length, but no more, of the corpuscle. The leech 
sucks up sporozoites with the blood and transfers them 
mechanically to another frog (tadpole). 

Dimensions. — Sporozoite 10-15 /a in length. 

Remarks. — ^The cycle as described above is based on the 
researches of Noller (1912, 1913, 1920). As Wenyon has 
remarked : " if this cycle of development is confirmed, it is 
a remarkable one in that the whole development, up to the 
formation of sporozoites, takes place in the endothehal ceUs 
of the blood-vessels, and is an illustration of a coccidium, 
originally transferred from host to host in the oocyst stage, 
as in the more typical forms, having become adapted to hfe 
in the blood-stream. . . . The possibihty of the escape of the 
oocysts to the exterior having been lost by this change of 
habit, the difficulty is overcome by the leech transferring 
from host to host." 

Some observers have described the vermicule as becoming 
spherical and undergoing schizogony in the red blood-corpuscles, 



but Noller believes that such schizonts belong to Dactylosoma 
ranarum, another parasite of the frog's corpuscles. 

Habitat. — ^Blood of Rana tigrina Daud., and in the leech 
which transmits it : Madras, Madras ; also from the same 
host from India in the Zoological Gardens, London. 

Fig. 72. — Lanhesterella minima (Chaussat). A, young schizont ; 
B, schizogony in an endothelial cell ; C, young forms in 
endothelial cell ; D, mierogametocyte ; E, formation of 
microgametes in the endothelial cells of blood-vessels ; 
F, fertilization of a female gamete in an endothelial cell of 
blood-vessel ; G, oocyst formed in an endothelial cell ; 
H, mature oocyst containing sporozoites in endothelial cell. 
(From Wenyon, after Noller.) 


94. Lankesterella monilis (Labbe). (Fig. 73.) 

Hsemogregarinaranarum (part), Celli & Sanfelice, 1891, p. 504, pi. v, 

figs. 2, 3, 4, 12-15 ; Kmse, 1890, p. 541. 
Drepanidiwm monile, Labbe, 1894, p. 76, pi. iii. 
Lankesterella monilis, Labbe, 1899, pp. 74-5, fig. 140. 
^Lankesterella monilis, Berestneff, 1903, pi. viii, fig. 7. 
Lankesterella monilis, Minchin, 1903, pp. 265, 267, 345. 

Free stages very mobile and showing successive undulations 
in locomotion. Body shows constriction into three segments, 
the middle one containing the nucleus. Nucleus vesicular, 

Fig. 73. — Lankesterella monilis (Labbe). 
Free forms. (After Labbe.) 

containing a karyosome and numerous chromatoid granules. 
No vacuole. Cysts as in L. ranarum. 

Habitat. — Blood of Eana tigrina Daud. and R. limnocharis 
Wiegmann : Bombay, Bombay. 

Subfamily CYCLOSPORINS Wenyon, 1926. 

Oocysts contain two sporocysts, each containing two, four, 
or eight sporozoites. 

Genus ISOSPORA Aime Schneider, 1881. 

Isospora, Aime Schneider, 1881, p. 401. 

Coccidium, Grassi, 1881, p. 135, pi. iii, figs. 37-40 ; p. 192. 

Diplospora, Labbe, 1893, p. 407. 

Hijaloklossia, Labbe, 1896, p. 535 ; 1899, p. 55. 

Isospora, Labbe, 1899, p. 72. 

Diplospora, Labbe, 1899, p. 71. 

Isospora, Laveran & Mesnil, 1902 c, p. 83; Castellani & Chalmers, 
1919, pp. 473, 475; Dobell, 1922, pp. 1497-8; Craig, 1926, 
pp. 349-53 ; Wenyon, 1926, pp. 806-28 ; Knowles, 1928, pp. 351- 
8 ; Reichenow, 1929, pp. 955-61 ; Kudo, 1931, pp. 274-5 ; 
Calkins, 1933, p. 565 ; Hoare, 1933, pp. 364-9, 383, 385. 

Oocyst develops two sporocysts, each containing four 

Remarks. — Species belonging to this genus are known from 
man, cats and dogs, birds, lizards, and frogs. Cats and dogs 


harbour three distinct species. One, or probably two, species 
are known from man. Becker (1934) gives the names and 
hosts of 42 named species and the hosts of 4 unnamed species 
oi Isospora. 

Key to Indian Species. 

1 (12). Infection limited to epithelial cells. .. 2. 

2 (4). Oocysts spherical 3. 

3. Oocysts 18-23|u, in diameter; sporo- 

cysts 12-15 fj. by 8-10 /u.. Intra- [Das-Gupta, p. 170. 

nuclear /. knowlesi Ray & 

4 (2). Oocysts not spherical 5. 

5 (10). Oocysts egg-shaped 6-9. 

6. Oocysts 25-33 ;u by 12-5-16 ^u ; sporo- 

cysts 12-14 ;u. by 7-9 ^i. In man ... /. belli Wenyon, p. 163. 

7. Oocysts 35-45 /x by 25-35 ;li; sporo- 

cysts 18-4 fi by 11-4 jLt. In cats ... J./eZ^s Wenyon, p 168. 

8. Measurements of oocysts and sporo- [p. 167. 

cysts not recorded. In lizards .... /. calotesi, sp. nov., 

9. Oocysts 15; 1-5 jx; sporocysts [Das-Gupta, p. 170. 

7'5 |Lt in diameter. In cobra /. minuta Mitra & 

10 (5). Oocysts subcylindrical 11. 

11. Oocysts 16-20 /x by ll-14ju.; sporo- [Das-Gupta, p. 172. 

cysts 8 ;u. by 4 ju. /. wenyoni Ray & 

12 (1). Infection may extend to the sub- 

epithelial tissue also 13. 

13 (14). Larger oocysts 18-20 /x by 14-16 ;Lt (in 

dogs); smaller oocysts 10-14 /x by 

7-5-9 yu, (in cats and dogs); sporo- [p. 165. 

cysts 13-5-15-5/x by 9-10 /x /. higemina (Stiles), 

14(13). Oocysts 20-25 /x by 15-22 /lx; sporo- ' [p. 171. 

cysts 16 /L4 by 10/x , . . . /. rivclta (Grassi), 

95. Isospora belli Wenyon. (Fig. 74.) 

Isospora liojninis, Dobell, 1922, pp. 1497-8, fig. 533 A. 
Isospora belli, Wenyon, 1923 a, p. 269. 
'fisospora belli, Knowles, 1924, p. 64. 
Isospora belli, Wenyon, 1926, pp. 820-4 fig. 350, 7-10; 1926 a, 

pp. 253-66. 
Isospora hominis, Dobell, 1926, pp. 74-85 ; Craig, 1926, pp. 349-53, 
fig. 62. 
'\Isospora belli, Knowles, 1928, pp. 356-8, fig. 81, 4, 17-21. 
Isospora belli, Reichenow, 1929, p. 958, fig. 923 ; Kudo, 1931, 

p. 274, fig. 114, a-c ; Calkins, 1933, p. 405. 
jlsospora belli, Knowles, 1933, p. 53 ; Das-Gupta, 1934, pp. 133-4, 
pi. ii. 

Only the oocysts and sporocysts are known. The former 
are elongate, egg-shaped, with one end more constricted 
than the other, forming a kind of neck. They are transparent, 
colourless bodies, with a wall consisting of two layers, the outer 
thick and porcellanous, the inner thin and membranous. 
At the narrow end there is an indication of a micropyle. 
In fresh stools are found unripe oocysts with cytoplasmic 




contents contracted into a spherical body with highly refractile 
granules, and a single nucleus seen as a clear pale area. The 
oocysts complete their development in one to four days, 
according to the temperature. The cytoplasm divides into 
two sporoblasts, which become elongated and covered with 
cyst- walls. In each sporocyst are developed four sporozoites 
and a large spherical residual body. Sporozoites are elongate 
structures, rounded at the anterior and tapering at the posterior 
end, and with a nucleus lying at the junction of the anterior 
and the middle third. 

The organism is probably a parasite of the epithehal cells 
of the small intestine, where schizogony and the development 
of the gametocytes will be found to occur. 

Fig. 74. — Isospora belli Wenyon. 

A, immature oocyst ; B, mature oocyst. 
(From Reichenow, after Dobell.) 

Dimensions. — Oocysts 25-33 ju. in length by 12-5-16 /x in 
width ; sporocysts 12-14 ju, by 7-9^. 

Pathogenicity.— Wenjon (1926) reports an observation by 
Conrad (1922) on a laboratory worker who accidentally in- 
gested material containing developed oocysts. Six days later 
diarrhoea set in and persisted for thirty days. The oocysts 
were found three weeks after the onset, and persisted in the 
patient's stools for 12 days, after which they disappeared, 
and recovery was complete. 

Similar symptoms and similar cysts have been found by 
a number of workers in different countries, but it is not certain 
if they belong to one or more species. 


Remarks. — The parasite named Isospora hominis (Railliet 
& Lucet, 1901) was first discovered by Virchow in 1860. 
Human Coccidia were observed, among others, by Woodcock 
(1915), Wenyon (1915), Cragg (1917), Dobell and O'Connor 
(1921), and Reichenow (1925), and referred to /. hominis, 
but according to Wenyon (1926) all these later findings of 
Isospora oocysts refer to I. belli and not to /. hominis, which 
he regards as a species with small oocysts. Dobell (1926) 
considers that the small subepitheHal form is identical with 
the larger form in the epitheUum. He holds that the case 
for two species is not proven, and that the name Isospora 
hominis should be adhered to. Wenyon (1926 a) has replied 
vigorously and adheres to his former opinion. Reichenow 
(1929) recognizes the two species as distinct. 

Cragg (1917) reported four cases from Bombay, but all these 
patients are beheved to have contracted the infection in the 
Mediterranean war area. Knowles (1928) reported having 
observed the infection in man five times during the six pre- 
ceding years, and reported another case in 1933. Das-Gupta 
(1934) recorded his observations on the case of a Bengah 
Brahman from Calcutta who had never been abroad. 

Habitat. — Faeces of man : Bombay, Bombay ; Bengal, 

96. Isospora bigemina (Stiles). (Fig. 75.) 

Corpuscles gemines, Finck, 1854. 

Cytospermmm villorum intestinalium cards et felis, Rivolta, 1874, 
-p. 1 ; 1877, pp. 42-6, 85-8. 

Coccidiwm bigeminum. Stiles, 1891, p. 163 ; 1892, pp. 517—26 ; 
Railliet & Lucet, 1891, p. 250 ; Labbe, 1896, p. 545 ; 1899, p. 67. 
'\Isospora bigemina, Castellani & Chalmers, 1919, p. 473. 

Isospora bigemina, Wenyon, 1923 a, p. 257, pi. xiii, figs. 1—11 ; 
Wenyon & Sheather, 1925, p. 10 ; Wenyon, 1926, pp. 809-13, 
figs. 343, 344; 1926, pp. 253-66; Knowles, 1928, pp. 351-5, 
fig. 81, 1; Reichenow, 1929, p. 957; Kudo, 1931, p. 274, 
fig. 114 d. 

Sporogony not confined to the epithelial cells ; it usually 
takes place and is completed in the subepitheHal tissue of the 
viUi. In the acute stages of infection reproduction occurs 
in the epitheU^im, and immature oocysts are passed in the 
faeces. Oocysts of two types. The smaller ones occur in 
both cats and dogs, while the larger ones have hitherto been 
seen only in dogs. The oocysts ripen in the gut-tissue rather 
than in the faeces : they have thin walls and the sporocysts 
may escape from them and be present in the faeces and carry 
the infection from one host to another. The sporocyst 
contains four sporozoites and a Httle residual substance in the 
form of a small clump or as dispersed granules. 

Dimensions. — Oocysts, smaller 8-14 /x by 7-9 fi, larger 
18-20 /Lt by 14-16ju. ; sporocysts 13-5-15-5)u, by 9-10/i (in dog). 



Remarks. — This parasite is usually found in the subepithelial 
tissue of the vilh, where it completes its sporogony, so that 
immature oocysts are not passed out in the fseces. Wenyon 
and Sheather (1925) had an opportunity of studying the 
intestine of a dog which had been killed durmg an acute phase 

Fig. 75. — Section of villus of cat, showing Isospora bigemina in the 
subepithelial tissues and Isospora felis in the epithelium 
( X 1000). Six mature oocysts of I. bigemina are seen in the 
subepithelial tissues. In the epithelium are seen a micro- 
gametocyte with microgametes, and one partially grown 
and two mature macrogametocytes. (After Wenyon.) 

of infection. The whole of the epithelium of the small intestine 
was found to be crowded with reproducing parasites ; but 
the subepithelial tissue was not invaded. The schizonts 
were small, measuring up to 5/x, and giving rise to eight minute 


merozoites . There were numerous macrogametocy tes measuring 
about 7-5 jLt in diameter. During life this dog passed in its 
faeces numerous immature oocysts measuring 10-14 /x. by 
7-5-9 )u,, the sporocysts measuring 7-5-9 ju, by 5-7 ju. In 
another puppy a large number of odcysts of the same dimensions 
were passed, and completed their development outside the 
body in the usual manner, forming two sporocysts, each with 
four sporozoites and a residual body. 

Wenyon (1926) is of the opinion that there are three species 
of Isospora in cats and dogs, and gives the dimensions of the 
oocysts as follows : — 

7. /eZis, 39-48 iLi by 26-37 /i. 

I. rivolta, 20-24 ^u by 15-20 ^it. 

/. bigemina (large), 18-20 /a by 14-16 jU,. 

I. bigemina (small), 10-16 ju. by 7-5-10 jU,. 

Dobell (1926) criticizes this view, but Wenyon (1926) 
adheres to it. Reichenow (1929) recognizes these species as 
distinct. According to Yakimoff, Matikaschwih, Rastegaieff, 
and Lewkowitsch (1930) the domestic cat harbours three 
species, viz., I. bigemina var. cati (Stiles), /. rivolta (Grassi), 
and I. felis Wenyon. Sen (1932) noted an unidentified 
coccidian from a dog at Muktesar. 

Habitat. — Subepithehal tissue of the intestine in dogs : 
Ceylon, Colombo. 

97. Isospora ealotesi, sp. no v. 

■fisospora sp., Setna, 1933, p. 97. 

Schizonts spherical. Multiple fragmentation gives rise to 
a number of small ovoid merozoites which gradually become 
vermiform. Merozoites vary in size : there may be four large 
merozoites clustered round a central cytoplasmic residue, 
or a large number, up to about 100, of slender fusiform mero- 
zoites. Male and female gametocytes may develop in the same 
epithelial cell or in separate ones, and in the earher stages are 
difficult to distinguish from the schizonts and from one 

Remarks. — The developmental stages are clearly marked off, 
and as many as 80 per cent of the hzards examined were 
infected. Infection was Hmited to the epithehum ; the sub- 
epithelial tissue was not affected. 

Dimensions.' — Schizonts up to 22 ;u, by 17 )u,; merozoites, 
small and slender 9)u, by 1-2 ^u,, large 8ju, by 2-8 ju,. 

Habitat.' — Intestine of the lizard, Calotes versicolor (Daudin) : 
Bombay, Bombay. 



98. Isospora felis Wenyon. (Figs. 75 & 76.) 

Isospora felis, Wenyon, 1923 a, p. 248, pis. ix-xii ; 1926, pp. 808, 

814, figs. 342, 344-8 ; 1926 a, pp. 253-66. 
■\Isospora sp., Knowles, 1928, p. 354. 
Isospora felis, Reichenow, 1929, p. 956, figs. 921 A, 922 ; Kudo, 

1931, pp. 274, fig. 114/. 
■fisospora sp. (" type B "), Knowles & Das-Gupta, 1934, pp. 387-90, 
pi. vii, figs. 1-4, 6. 

Development takes place in the epithelial cells only of the 
small intestine and not in the deeper layers of the villi. Oocysts 
egg-shaped, with one pole somewhat narrowed. The oocysts 

Fig. 76. — Mature oocyst oi Isospora felis Wenyon. 
(From Reichenow, after Dobell.) 

drop out of the epitheHal cells into the lumen of the gut 
and pass out with the faeces, the formation of sporocysts and 
sporozoites taking place in the oocysts while outside the body 
of the host. The macrogametocyte is easily distinguished 
in shghtly advanced stages, as it has a single nucleus with 
a characteristic appearance. In the microgametocyte the 
nucleus divides repeatedly and hundreds of microgametes 
develop in each. Each microgamete is a flexible rod-like 
body with two flagella. FertiHzation takes place while the 
macrogamete is within an epithelial cell. The oocyst develops 


a thick chitinous wall after it has passed into the lumen of the 
intestine. Oocysts containing two sporoblasts may be found 
in the gut, and in this condition are passed out in the faeces. 
Outside the body the sporoblasts in the oocyst elongate, acquire 
a double contour, and become sporocysts. In each sporocyst 
are developed four club-shaped sporozoites and a large rounded, 
granular, residual body. 

Dimensions. — Oocysts, egg-shaped type 35-45^ by 25-35 )u. 
(39-48 |U by 26-37 ^u, according to Wenyon), rounded type 
25-37 /i by 21-37 ^u,; microgamete 3-1 ft; sporocysts 18-4/x 
by 11-4^. 

Remarks. — ^Knowles (1928) stated that Isospora infection 
was not micommon in cats in Calcutta, and that infection 
was absolutely hmited to the epitheHal cells, but did not mention 
the name of the species or give the dimensions of the oocysts. 
On my request for further information, Knowles and Das- 
Gupta (1934) examined the stools of 13 cats and 8 kittens, 
and by the employment of concentration method found 
14 out of 21 animals examined to be infected. In all they 
measured 353 oocysts. After discussing the views of Wenyon 
and of Dobell they are of opinion that the evidence collected 
by them seems to point to two " types " being present, 
though they are not in a position to assert that they are 
different species. The smaller, oval type of oocyst encountered 
measured from 20-4 ju, by 15-3 ju, to 47-6//, by 40-8 ju,, and the 
larger, egg-shaped type of oocysts measure 38-45 [x by 
27-36 /x. 

Their " type A" or small oocysts are always perfectly oval 
and resemble /. rivolta, and the " tj^^pe B " or large oocysts 
are more egg-shaped, pyriform or ovoid, and resemble I. felis. 
I am inchned to think that the form of the oocyst and the 
measurements given show that the two " types " are distinct 
species, as is held by Wenyon and Reichenow. 

The vast majority of the oocysts seen by Knowles and Das- 
Gupta in freshly passed faeces were in the unsegmented state, 
with the protoplasmic contents present as a single spherical 
mass within the thick oocyst waU, while in a few the first 
nuclear division had taken place and two sporoblasts were 
forming, although the sporocysts had not yet formed. They 
also noted the occurrence of a fair number of motile sporozoites 
in the contents of the jejunum and ileum, both in the fresh 
preparations and in fixed and stained films, from which 
they conclude that an occasional oocyst may develop to maturity 
within the lumen of the gut, although the vast majority are 
passed in the unsegmented state. 

Habitat. — ^Epithelial cells of the small intestine of cat, 
Felis domesticus Linn. : Bengal, Calcutta. 



99. Isospora knowlesi Ray & Das-Gupta. (Fig. 77.) 

■fisospora sp., Knowles & Das Gupta, 1935, pp. 703, 705, pi. xxx, 

figs. 14, 20. 
Isospora knowlesi, Ray & Das-Gupta, 1937 c, pp. 269—74, pi. vii. 

Young gametocyte intranuclear. Male gametocyte gives 
rise to biflageUate gametes which drop into the lumen of the 
intestine or penetrate the neighbouring epithehal cells. 
Oocysts thick-walled, spherical. Sporocysts elhpsoidal, with 
a knob-Hke structure at one pole. Sporozoites arranged 
regularly, with large sporocystic residue. Unsegmented 
or segmented but immature oocysts discharged from the host. 
Sporulation takes place in 4 to 5 days. 

Fig. 77. — Isospora knowlesi Ray & Das-Gupta. A, oocyst within the 
nucleus of a host-cell ; B, oocyst from the faecal matter ; 
C, manure oocyst. (After Ray & Das-Gupta.) 

Dimensions. — Oocysts 18-23 /x in diameter; sporocysts 12- 
15^ by 8-10/x. 

Remarks. — The oocysts resemble those of Isosjiora mesnili 
Sergent (1902) in being intranuclear in habitat, and approach 
those of/, camilleri Hagenmuller (1898) in size. 

Habitat. — Nuclei of epithelial cells oiF the small intestine of 
Hemidactylus flaviviridis (Riippell) : Bengal, Calcutta. 

100. Isospora minuta Mitra & Das-Gupta. 

Isospora minuta, Mitra & Das-Gupta, 1937 a, p. 291. 

Oocysts disporocystid, final stages of sporogony taking place 
outside the body of the host. Sporocysts spherical. Oocystic 
residue absent. 

Dimensions. — Maximum size of oocyst 15 /a by 7-5 /x ; sporo- 
cyst 7-5 /A in diameter. 

Remarks. — Full description of the species has not yet been 


published. The form is said to differ from /. dirumpens 
Hoare in that development inside the oocyst takes place 
outside the body of the host. The disporocystid condition of 
the oocyst was evidenced after being kept in 1 per cent, chromic 
acid for three days. It also differed from /. naiee Fantham 
in that there was no oocystic residuum. 

Habitat. — ^In the fsecal matter oiNaja naja (Linn.) : Bengal, 

101. Isospora rivolta (Grassi). (Fig. 78.) 

Goccidium rivolta, Grassi, 1879, p. 135, pi. xxxiii, figs. 41-4 ; 1881 a, 

p. 632. 
Goccidium bigeniinuni, Labbe, 1899, p. 67. 

Isospora rivolta. Brown & Stammers, 1922, p. 1165; Wenyon, 
1923 a, p. 260, figs. 12-15 ; 1926, pp. 808, 813, figs. 342, 345, 
346, 350. 
■fisospora sp., ("type A "), Knowles, 1928, p. 354, fig. 81, 2. 
Isospora rivolta, Reichenow, 1929, pp. 956-7, fig. 921 b ; Kudo, 1931, 

p. 274, fig. 114e. 
■fisospora rivolta, Knowles & Das-Gupta, 1931, pp. 175-6, pi. ix. 
■\Isospora sp. ("type A"), Knowles & Das-Gupta, 1934, pp. 387-90, 

pi. vii, figs. 5, 6. 
"fisospora rivolta, Aiyar, 1937, p. 402. 

Oocysts ovoid, with a double contour. In each are developed 
two sporocysts, and a small residual body may be present. 
The sporocysts are elongate bodies with rounded ends. 

Fig. 78. — Mature oocyst oi Isospora rivolta (Grassi). 
(After Reichenow.) 

Each sporocyst contains four sporozoites and a large residual 
body filled with globules of a refractile material. The infection 
is usually confined to the epithehal layer, but the form may 
also reproduce in subepithehal tissues of the alimentary canal 
of the host and produce male and female gametocytes. 

Dimensions. — Oocysts 20-25 /x in length by 15-22^ in 
breadth ; sporocysts 16 ju, by 10|Li. 



Remarks. — This is a common parasite of cats and dogs. 
Knowles and Das-Gupta (1934) have recorded the measurements 
of 253 oocysts and consider the smaller oocysts, which are 
perfectly oval and measure 23-28 ju, by 18-23 /x, as belonging 
to " type A," which corresponds to I. rivolta (Grassi). 

Knowles and Das-Gupta (1931) also found in the csecal 
contents of a small Indian mongoose immature oocysts of 
a coccidium containing a single rounded unsegmented mass of 
protoplasm, and studied its development in a moist-chamber. 
The average dimensions of the oocyst were 20-6 ju, by 17-2^. 

Habitat. — Epithehal cells of the small intestine of cat, Felis 
domesticus Linn. : Bengal, Calcutta ; also in the caecal 
contents of the mongoose, Herpestes auropunctatns (Hodgs.) : 
Bengal, Calcutta. 

102. Isospora wenyoni Ray & Das-Gupta. (Fig. 79.) 

■fisospora wenyoni, Ray & Das-Gupta, 1935 a, pp. 219-24, pi. viii. 
Young intracellular trophozoites are found in the epithehal 

Fig. 79. — Isospora wenyoni Ray & Das-Gupta. A, merozoite entering 
an epithelial cell ; B, merozoites escaping into the lumen 
of the intestine ; C, microgametocyte ; D, microgametes 
clustered round a central mass of cytoplasm ; E, macro- 
gametocyte showing spherical nucleus and the karyosome ; 

F, macrogamete showing a recurved tail, an elongated 
nucleus, and two microgametes near the posterior end ; 

G, mature oocyst. (After Ray and Das-Gupta.) 


cells of the small intestine. Young schizonts possess eight to 
twelve nuclei, and schizogony results in producing the same 
number of merozoites. Each merozoite is spindle-shaped, 
and carries a pair of hyaline blades or laminae at its anterior end, 
which help it to enter the epithehal cell. The male gametocyte 
shows a large niunber of nuclei situated towards its periphery, 
and the mature microgametes are found to cluster round 
a residual mass of cytoplasm. The macrogametocyte is 
distinguished by its size and by the presence of darkly staining 
granules in its cytoplasm ; its posterior end shows a short 
recurved tail ; the nucleus at an early stage is spherical and 
contains a karyosome, but when mature becomes elongated, 
and the karyosome breaks up into a number of small, irregularly 
scattered granules. Oocysts are subcyhndrical, disporocystid, 
and tetrazoic, and develop outside the host. There is no 
oocystic residuum, and the oocystic membrane has a double 
contour. Sporocysts have their long axis directed at right 
angles to the long axis of the oocysts ; a sporocystic residuum 
is present. 

Dimensions. — ^Trophozoites, young 10 /x by 3 /x ; schizonts, 
young 20-25 ;u,; merozoites 12ju, by Sju, ; microgametes 
2-4)u, by 1-5 /x ; macrogametes 16-20 /x by ll-14)u. ; oocysts 
16-20 ju. by 11-14/x ; sporocysts 8/x by 4ju.. 

Habitat. — In the small intestine of the toad, Bufo melano- 
stictus Schneider : Bengal, Calcutta. 

103. Isospora sp. 

"flsospora sp., Cooper & Gulati, 1926, pp. 191-2. 

Oocysts of an unidentified species of Isospora were found in 
the faeces of a heifer calf at Dhupatal, of a bull calf and two 
cows at Tocklai, and of a heifer at Shillong, in Assam. 

Oocysts were quite round, and within twenty-four hours 
two fully formed sporoblasts were seen in all cases. 

Habitat. — ^Ahmentary canal of cows : Assam, Dhupatal, 
Tocklai, and Shillong. 

Subfamily EIMERIIN.E Wenyon, 1926. 

Oocysts contain four sporocysts, each containing two, four, 
or many sporozoites. 

Genus EIMERIA Aime Schneider, 1875. 

Gregarina, Eimer, 1870, p. 4. 
Eimeria, Aime Schneider, 1875 a, pp. xl-xlv. 
Psorospermium, Rivolta, 1878. 
Coccidium, Leuckart, 1879, p. 254. 
Orthospora, Aime Schneider, 1881, p. 389. 


Crystallospora, Labbe, 1896, p. 554 ; 1899, p. 63. 

Goussia, Labbe, 1896, p. 551 ; 1899, pp. 63-5. 

Eimeria, Labbe, 1899, pp. 58-9. 

Paracoccidium, Laveran & Mesnil, 1902 6, p. 858. 

Eimeria, Castellani & Chalmers, 1919, pp. 475-6 ; Dobell, 1919 a, 
pp. 147-97 ; Snijders, 1920, pp. 427-32 ; Dobell, 1922, pp. 1498-9. 

Jarrina, Leger & Hesse, 1922, pp. 74-7. 

Eimeria, Craig, 1926, pp. 353-8 ; Thomson & Robertson, 1926, 
pp. 282-3; 1926 a, pp. 420-1; Wenyon, 1926, pp. 828-62; 
Knowles, 1928, pp. 358-66 ; Reichenow, 1929, pp. 937-55 ; 
Kudo, 1931, pp. 271-4 ; Stiles, 1929, p. 881 ; Levine & Becker, 
1933, pp. 83-106 ; Calkins, 1933, pp. 403, 565 ; Hoare, 1933, 
pp. 374-7, 378, 381, 383, 385 ; Wenrich, 1935, pp. 7-9 ; Ray & 
Das-Gupta, 1936, p. 345-6 ; 1937 d, pp. 275-7. 

Oocyst develops four sporocysts, each containing two 

Remarks. — ^The typical life-history is as described by 
Schaudinn for E. schubergi, of which a brief summary is given 
for ready reference. Oocysts gain entrance into the host 
through the mouth. Sporozoites escape from the sporocysts 
and, passing through the micropyle of the oocyst, move about 
in the lumen of the gut till they enter the epithelial cells 
of the gut- wall, where they grow into schizonts. Schizonts 
are large rounded bodies, which give rise to merozoites. 
The latter escape into the lumen of the gut and enter new 
host-cells and repeat the process. Some merozoites develop 
into macro- and microgametocytes. The macrogametocyte 
produces a single macrogamete after extruding part of its 
nuclear material. The microgametocyte produces a number 
of biflagellate microgametes. Sjmgamy takes place and the 
zygote secretes a membrane around itself, forming an oocyst. 
The nucleus divides twice and four sporoblasts are developed 
inside the oocyst. Each sporoblast secretes a membrane 
and becomes a sporocyst, and two sporozoites are developed 
inside each sporocyst. Oocysts pass out in the faecal matter 
of the host and infect other hosts by being ingested. 

The genus has a very wide distribution, numerous species 
having been described from all classes of Vertebrates, terrestrial, 
freshwater, and marine, and a few Invertebrates" (myriopods, 
etc.). They have been found to occur as intestinal parasites 
in man, horses, cattle, pigs, sheep and goats, rats and rabbits 
domestic birds such as fowls, ducks, pigeons, pheasants, etc. 
a few lizards and tortoises ; in frogs, newts, and salamanders 
in fish ; in centipedes, etc. According to Levine and Becker 
(1933) no less than 220 species of this genus have been recorded 
from 183 species of hosts of widely different groups. These 
hosts include Annelids, 2 species ; Myriapods, 6 species ; 
Insects, 3 species ; Enteropneusta, 1 species ; Fishes, 43 
species ; Amphibia, 10 species ; Reptiles, 26 species ; Birds, 
24 species ; and Mammals, 60 species. The shape, dimensions, 



and colour of the oocyst, the appearance of the oocyst wall, 
the character of the micropyle, and the presence or absence 
of a residual mass when the sporoblasts separate, as also the 
shape and size of the sporocyst, are taken into account in 
identifying the species. Allen (1934) gives a key to twenty- 
four known species of Eimeria in birds. 

According to Dobell (1932), Leeuwenhoek probably first saw 
the oocysts of rabbit Coccidia as far back as 1674, but, as 
remarked by Wenrich (1935), it has only recently been demon- 
strated beyond a doubt that rabbit Coccidia are peculiar to 

Fig. 80. — Life-cycle of Eimeria schubergi (Schaudinn). (x400.) 
A, entrance of a sporozoite in the gut epithelium of the host 
and growth of schizont ; B, three stages in schizogony to 
form merozoites, which repeat schizogony ; or G, become 
macro- and microgametocyte ; D, E, formation of macro- 
gamete ; F-H, formation of microgaixietes ; I, mature 
gametes and fertilization ; J, secretion of a membrane round 
the zygote ; K—N, stages in sporocyst formation ; 0, oocyst 
containing foxir sporocysts, each with two sporozoites ; 
P, escape of the sporozoites. (From Kudo, after Schaudinn.) 

rabbits and difierent from those of cattle and poultry ; and 
further, that rabbits may harbour five or six different species 
of Eimeria, of which E. stiedss, occurring in the Hver, is the one 
most often causing disease and death ; the other species 
live in the intestine. 


Key to Indian Species. 

1 (15). Oocysts spherical 2. 

2 (8). Sporocysts with rounded ends 3. 

3. Oocysts 18-21 |Li in diameter, without 

residium ; sporocysts 10^ by 7 fx, [han), p. 177. 

with residuum. In fish and man . . E. clupearum (Thelo- 

4. Oocysts almost spherical (sometimes 

oval), 16-20^ by 14-18 /x, without 

residuum ; sporocysts spherical or [p. 184. 

oval. In lizards E. knowlesi Bhatia, 

5. Oocysts 16-18 /x in diameter, without 

residuum; sporocysts oval, with [p. 186. 

residuum. In tortoises E. legeri (Simond), 

6. Oocysts 20-22 /x, without residuum ; 

sporocysts oval, with residuum. In [(Simond), p. 183. 

gharials E. kermorganti 

7. Oocysts spherical or almost spherical, 

12-25 ;i in diameter; sporocysts 

pear-shaped, 9-9-11/x by 5-3-5-7/x; [p. 194. 

without residuum. In cattle E. zurni (Rivolta), 

8 (2). Sporocysts fusiform or with pro- 

jections 9. 

9 (11). Sporocysts elliptical, narrow end pro- 

jecting as a neck and bearing an 

inverted V-shaped appendage 10. 

10. Oocysts 9-13 /x in diameter; sporo- [Bana, p. 181. 

cysts 9 /x by 4 j(x. In fish E. harpodoni Setna & 

11 (9). Sporocysts fusiform 12. 

12. Oocysts 36-52 /x in diameter, with 

residuum ; sporocysts fusiform, 

30-32 /x by 7*5 /x, with residuum. In [p. 189. 

fish and man E. sardinse (Thelohan), 

13. Oocysts 14/x in diameter, without resi- 

duum ; sporocysts spindle-shaped, [p. 184. 

with residuimi. In tortoise E. koormse Das-Gupta, 

14. Oocysts 8-11 ;ix in diameter, no resi- 

duum; sporocysts 4'5-6fx by 3/x; 

schizonts bear laminae at anterior [p. 185. 

end. In toads E. Imninata Ray, 

15 (1). Oocysts not spherical 16. 

16 (18). Oocysts mitre-shaped 17. 

17. Oocysts with 4 or 5 projections, 10- 

15/x in diameter, without residuum ; [Mesnil), p. 187. 

sporocysts oval. In tortoises E. mitraria (Laveran & 

18 (16). Oocysts subspherical, ovoid, oval or 

cylindrical 19. 

19 (24). Oocysts subspherical, ovoid or lemon- 

shaped 20. 

20 (21). Oocysts subspherical, 16-4 ;« by 14-35 /x. 

Oocystic residuTom present. In [Das-Gupta, p. 178. 

pigeons E. columbae Mitra & 

21 (20). Oocysts ovoid or lemon-shaped 22. 

22(23). Oocysts lemon-shaped, 17-20-4 |u by [& Das-Gupta, p. 182. 

13-6-17 /LA. In lizards E. hemidactyli Knowles 

23 (22). Oocysts usually ovoid, 20-40 /ix by 

17— 26ju,, sometimes spherical, 18/xin 

diameter; sporocysts 13 /x by 6 /x, [Marotel), p. 179. 

with residuum. In sheep E.faurei (Moussu & 


24 (19). Oocysts oval or cylindrical 25. 

25 (34). Oocysts oval 26. 

26 (31). Sporocysts ovoid or oval 27. 

27. Oocysts 17-34/i by ll-15;(i, without 

residuum ; sporocysts ovoid, 7-9 ^ 

by 5-1 II, with residuum. In [Bana, p. 179. 

lizards E. flaviviridis Setna & 

"28. Oocysts egg-shaped, 25-35 /x, without 

residuum ; sporocysts pear-shaped, 

10-8-14-4/Li by 7-9^, with residuum. [& Galouzo, p. 191. 

In cattle E. smithi YakimoS 

29. Oocysts 32-4-41-4;Li by 21-6-28-8/i, 

without residuum ; sporocysts oval, 

13-19 /x by 3-5-4: fi, without resi- [gaieff, p. 194. 

duum. In nilgau E. yahimovi Raste- 

30. Oocysts egg-shaped, 18ju, by 14-4|U,. [gaieff, p. 194. 

Sporocysts not known. In deer. . . E. wassilewskyi Raste- 

31 (26). Sporocysts fusiform 32. 

32(33). Oocysts 18-8-27-04/i by 18-8-20-8/x, 

without residuum ; sporocysts 14-6 /x [Gupta, p. 188. 

by 6 jLt. In snakes E. najse Ray & Das- 

33 (32). Oocysts 29-31 /x by 22-5-24-5/x, with 

residuum ; sporocysts 14 /x by 4-6 /x. [Gupta, p. 189. 

In snakes E. piscatori Ray & Das- 

34 (25). Oocysts cylindrical 35. 

35 (36). Oocysts 36 ;u, by 18 /x, with residuum. [Das-Gupta, p. 179. 

In snakes E. oylindrica Ray & 

36 (35). Oocysts 25 /x by 12 /x, with residuum ; 

sporocysts oval, 10-12 /x in length. [p. 193. 

In fish E. southwelli Halawani, 

104. Eimeria clupearum (Thelohan). (Fig. 81.) 

Coccidium sp., Thelohan, 1892, p. 158, pi. xii, figs. 13, 14. 
Coccidium clupearum, Thelohan, 1894, p. 565, pi. xxii, figs. 19, 20. 
Goussia clupearum, Labbe, 1896, p. 552, pi. xviii, figs. 24, 25; 

1899, p. 64. 
Eimeria sp., Wenyon, 1915 b, p. 1404. 
Eimeria wenyoni, Dobell, 1919 a, pp. 187-8, pi. viii, fig. 2 ; Castel- 

lani & Chahners, 1919, pp. 475-6 ; Dobell & O'Connor, 1921, 

p. 100, pi. vi, fig. 104. 
■\Eimeria clupearum, Knowles, 1924, p. 24 ; 1928, p. 362, fig. 83. 
Eimeria clupearum, Wenyon, 1926, pp. 851-2, 861, fig. 350, 11 ; 

Thomson & Robertson, 1928, pp. 360-1, 363; fig. 83, 2; 

Reichenow, 1929, pp. 951, 952-3, fig. 918 b ; Kudo, 1931, p. 274, 

fig. 113/1. 

The oocyst is spherical, is of a hght brown colour, and possesses 
a fairly thick wall, roughened on the outer surface and lined 
by a dehcate membrane on the inner surface. It contains 
four sporocysts, each having rounded ends. Each sporocyst 
contaias two sporozoites and some residual substance in the 
form of one or two masses of refractile material. Each sporo- 
zoite, with rounded anterior and pointed posterior end, con- 
tains an ovoid refractile body in its anterior portion. 

Dimensions. — Oocyst 18-21 /a in diameter; sporocyst 10 /a 

SPOR. 21 



Remarks. — ^This form was originally described as a parasite 
of the Hver of herrings, mackerel and sprats. Wenyon 
(1915 h) found the ripe oocysts in the faeces of a patient 
retm-ned to England from GaUipoH. Dobell (1919 a), con- 
sidering it to be a human parasite, named it E. wenyoni. 

Fig. 81. 

Fig. 8i 

Fig. 81. — Oocyst of Eimeria dupearum, (Thelohan). (After Dobell and 

Fig. 82. — Eimeria faurei (Moussu & Marotel). A, immature oocyst ; 

B, mature oocyst. (From Wenyon, after Noller, Schur- 

johann, and Verbrodt.) 

Knowles (1924) recorded it from the fseces of a man at Calcutta. 
Thomson and Robertson (1926) have shown that the oocysts 
ofE. dupearum occur in large numbers in the liver of herrings, 
mackerel and sprats, and that tliis is the source of the 
oocysts in human fseces. 

Habitat. — ^Fseces of man : Bengal, Calcutta. 

105. Eimeria columbse Mitra & Das-Gupta. 

'\Eimeria columbse, Mitra & Das-Gupta, 1937 6, p. 291. 

Oocysts subspherical, tetrasporocystid. Oocystic residuum 
present. Sporocyst dizoic. 

Dimensions. — ^Maximum size of oocyst 16'4/x by 14'35/x. 

Remarks. — Full description of the species has not yet been 
pubhshed. Tetrasporocystid dizoic condition was observed 
after keeping the oocysts in 1 per cent, chromic acid for four 
days. It is said to differ from E. avium (Silvestr. & Rivolta) 
in the oocysts being subspherical, and from E. pfeijferi Labbe 
in oocystic residuum being present. 

Habitat. — Intestine of the pigeon, Columba livia intermedia 
Strickland ; Bengal, Calcutta. 


106. Eimeria cylindrica Ray & Das-Gupta. 

'\Eimeria cylindrica, Ray & Das-Gupta, 1936 a, p. 345. 

Oocysts cylindrical, measuring 36 ju. by 18 /u,. Oocystic 
residuum present. 

Habitat. — Rectum of snake, Natrix piscator (Schneid.) : 
Bengal, Calcutta. 

107. Eimeria faurei (Moussu & Marotel). (Fig. 82.) 

Coccidium sp., Moussu & Marotel, 1901, pp. 1087-9. 

Goccidium faurei, Moussu & Marotel, 1902, pp. 82-98, 10 figs. 
"f Coccidium faurei, Baldrey, 1906, p. 387. 

Eimeria faurei, Noller, Schurjohann. & Verbrodt, 1922, p. 993,. 
7 figs. ; Wenyon, 1926, pp. 843-4, fig. 363 ; Reichenow, 1929, 
pp. 945-6 ; Rastegaieff, 1930, pp. 379, 399 ; Kudo, 1931, p. 272. 

Oocysts usually ovoid, but sometimes spherical, and possess 
a definite micropyle closed by a cap. Four sporo blasts are 
developed in the oocyst, and a residual body may or may not 
be present. Sporocysts possess a micropyle at the more 
pointed end ; a residual body is always left in the sporocyst. 

Dimensions. — Oocyst, ovoid 20-40 ^u, by 17-26 ^a; spherical 
18jLt in diameter ; sporocysts ISfxhy Qfx 

Eemarks. — The parasite is said to cause progressive pernicious 
anaemia that sooner or later ends fatally. 

Habitat. — Intestine of sheep : India (locahty not noted). 

108. Eimeria flaviviridis Setna & Bana. (Fig. 83.) 

■fEimeria sp., Setna, 1933, p. 97, fig. 2 ; Ray & Das-Gupta, 1935 6,. 

p. 315. 
fEimeria (" species B "), Knowles & Das-Gupta, 1935, pp. 703, 705, 

pi. XXX, figs. 6—9, 17. 
fEimeria flaviviridis, Setna & Bana, 1935 b , pp. 256-60, pis. i-iii; 

Ray & Das-Gupta, 1937 c, p. 270. 

Various developmental stages occur either in the cells of 
the epithehal fining of the gaU-bladder or attached to them, 
or floating freely in the fluid in the gaU-bladder. Schizonts 
rounded, oval, or irregular in shape and give rise to from 16 to 
140 merozoites. Merozoites are elongate curved bodies, 
with pointed extremities. Microgametocyte is a large ovoid 
body. Microgametes simple, rod-shaped. Macrogametocyte 
characterized by a crescentic mass of granules surrounding 
its nucleus. Oocysts colourless, elfiptical. Four sporoblasts 
develop as roundish masses, without the formation of any 
residual body. Sporocysts ovoid, without a cap : each con- 
sists of two valves, appfied to one another like two watch- 
glasses, and contains two sporozoites, which are elongated 
structures sfightly longer than the sporocyst, with their ends 
coiled in. The sporocystal residue is represented by a mass 
of granules lying in the middle. Development of the sporo - 




Fig. 83. — Eimeria flaviviridis Setna & Bana. A, section of a portion of 
the gall-bladder showing developmental stages in the epithe - 
lial cells {a, schizont; b, merozoites; c, macrogametocyte) ; 

B, schizont, showing repeated division of the nucleus ; 

C, formation of merozoites, with a large residual body in 
the middle ; D, macrogametocyte ; E, microgametocyte ; 
F, oocyst with four sporoblasts ; O, mature oocyst containing 
sporocysts, each with two sporozoites and a residual body. 
(After Setna and Bana.) 


zoites takes place while the oocysts and sporocysts are floating 
in the fluid in the gall-bladder. Oocysts pass into the alimentary 
canal and are expelled with the faeces, fresh infection taking place 
by these being ingested. 

Dimensions — Schizonts 12-21 //,; merozoites 8-10 )(a by 
1-3-1-5/i ; microgametocytes 16-17 /x ; oocyst 17-34 /x in length 
by 11-15/x in width ; sporocysts 7-9 /u, by 5-7^ ; sporozoites 
9-13/x by l-04;Li 

RemarTcs. — Setna and Bana (1935 h) described this parasite 
as a new species under the name of E.flaviviridis,3ind observed 
both schizogony and sporogony taking place within the gaU- 
bladder. Development of the oocyst is completed in 62 hours 
within the gall-bladder, or within 54 to 61 hours outside the 
body of the host under experimental conditions. 

Rarely oocysts containing three sporoblasts and three sporo- 
cysts were encountered, but one of these was double the size 
of the others and developed four sporozoites, the total number 
of sporozoites being thus the normal mmaber eight. 

Knowles and Das- Gupta (1935) have recorded three species 
from the same host. In their " species B " the oocysts range 
in size from 17-22^ in length by 12-15 /x in breadth. They 
also met with occasional giant forms measuring over 30 ju, 
in length, and remarked that they possibly corresponded 
with the species observed by Bana and reported by Setna 
(1933). The oocysts reported by Setna and Bana measured 
25-34 /x in length and 11-14^ in breadth. It is thus highly 
probable that E. flaviviridis described by Setna and Bana 
from Bombay and " species B " described by Knowles and 
Das-Gupta from Calcutta are identical. 

The species shows a very close resemblance to E. agamse 
Laveran & Pettit, 1910, as regards the occurrence of both 
schizogony and sporogony in the gall-bladder and bile-ducts 
and in the form and dimensions of the oocysts, but differs as 
regards the form of the sporocysts. In E. agamse the oocysts 
are more or less elongated oval, measuring 20-25 /x by 11-14)li, 
and the sporocysts fusiform, measuring 8/x by 4ju,. In 
E. flaviviridis the sporocysts are ovoid, and measure 7-9 /x 
by 5-7^. 

Habitat. — Gall-bladder and intestine of the Hzard, Hemi- 
dactylus flaviviridis (Riippell) : Bengal, Calcutta ; Bombay, 

109. Eimeria harpodoni Setna & Bana. (Fig. 84.) 

■fEimeria sp., Setna, 1933, p. 97, fig. 1. 

\Eimeria harpodoni, Setna & Bana, 1935 a, pp. 165-9, figs. 1-4. 

Oocysts almost spherical, with four sporoblasts and a large 
honeycomb-like residual body. Sporocyst elliptical in outhne,. 



with the narrow end of the oval projecting in the form of 
a neck, and bearing a broad, inverted, V-shaped appendage, 
clearly visible in both living and stained preparations. Each 
sporocyst contains two sporozoites, and a sporocyst residue is 
usually present at one end. Sporozoites usually slightly 
curved, with one end broader than the other. 

Pig. 84. — Eimeria harpodoni Setna & Bana. A, spherical oocyst with 
four sporoblasts, from the lumen of the gut ; B, mature 
oocyst with four peculiar-shaped sporocysts each containing 
two sporozoites and residue, after development outside the 
body. (After Setna and Bana.) 

Dimensions. — Oocysts 9-13 /x in diameter; sporocysts 
9 fxhy 4 fjL. 

Habitat. — Alimentary canal of the fish popularly known 
as " Bombay Duck," Harpodon nehereus (Ham.-Buch.) : 
Bombay, Bombay. 

110. Eimeria hemidactyli Knowles & Das-Gupta. (Fig. 85.) 

■\Eimeria hemidactyli, Knowles & Das-Gupta, 1935, pp. 703, 705, 
pi. XXX, figs. 10-13, 18, 19 ; Ray & Das-Gupta, 1937 c, p. 270. 

Oocysts lemon-shaped. No micropyle could be seen, but 
the wall of the oocyst was definitely thinner at the narrow 
anterior pole than at the posterior end. 

Fig. 85. — Eimeria hemidactyli Knowles & Das-Gupta. 
Lemon-shaped oocysts. (After Knowles and Das-Gupta.) 



Dimensions. — Oocysts 17-20-4/x by 13-6-17 /u.. 
Habitat. — Gut-contents oi Hemidactylusflaviviridis (Riippell): 
Bengal, Calcutta. 

111. Eimeria kermorganti (Simond). (Fig. 86.) 

"fCoccidiuni kermorganti, Simond, 1901 c, pp. 483-5, figs. 1—6. 
Eimeria kermorganti, Wenyon, 1926, p. 860 ; Reichenow, 1929, 
p. 949. 

In the young condition the macrogamete is a small, naked, 
granular sphere, provided with a nucleus and situated in a host- 
cell. The sphere grows till it reaches a diameter of 20-22 /*. 
After undergoing syngamy it secretes a double membrane 
round itself. The granular mass contracts and divides 
into four sporoblasts. Oocyst spherical ; contains four oval 

Fig. 86. — Eimeria kermorganti (Simond). A, growing macrogamete ; 
B, encysted condition ; C, oocyst containing sporoblasts ; 
D, oocyst containing four sporocysts ; E, fully developed 
sporocyst ; F, schizont dividing into merozoites. (After 

sporocysts without any residuum. Sporocyst oval ; con- 
tains two comma-shaped sporozoites, the larger ends of which 
are situated at opposite poles, and there is a small mass of 
sporocystal residuum. Schizogony was found in some cells, 
resulting in a large number of merozoites. 

Remarks. — It is rare to find a Coccidium in an organ from 
which there is no exit for the ripe oocysts. Simond beheved 
that the parasite must be occurring also in other organs con- 
nected with the intestine, but could not find them there. 

Habitat. — Spleen of the gharial, Gavialis gangeticus (GmeHn), 
of the River Ganges. 


112. Eimeria knowlesi Bhatia. (Fig. 87.) 

"fEimeria ("species A"), Knowles & Das-Gupta, 1935, p. 701, 

pi. XXX, figs. 1-5, 15, 16. 
Eimeria knowlesi, Bhatia, 1936, p. 177. 
"fEimeria (" species A "), Ray & Das-Gupta, 1937 c, p. 270. 

Oocysts almost spherical, sometimes oval. The mature 
oocyst contains four sporocysts, each containing two sporo- 
zoites, and does not show any micropyle. Sporocysts show 
no cap. 

Dimensions. — Oocysts from 16-20 /u, by 14-18 /x. 

Fig. 87.- — -Eimeria knowlesi Bhatia. A, spherical oocyst ; 
B, oval oocyst. (After Knowles and Das-Gupta.) 


Bemarks. — ^Knowles and Das-Gupta (1935) remark that the 
form may be Eimeria raillieti Leger, but it was not possible 
for them to make certain as they were unable to obtain 
Leger's paper in India, nor could it be procured even in 
London. This was due to the reference having been wrongly 
cited by Wenyon. Coccidium raillieti was originally described 
by Leger (C. R. Soc. Biol, xi, i (1899), pp. 309-11) from 
Anguis fragilis Linn., and an abstract of the paper is given 
in Wiegmann's ' Archiv ' (1904, pt. 3, pp. 115-16). The oocysts 
ofE. knowlesi correspond to those of ^. raillieti in size, but those 
of the latter are oval and show a small button-like protuber- 
ance at one pole. As, however, the oocysts in E. knowlesi 
are sometimes oval, and the size-range is not markedly 
different from that of E. flaviviridis ( = " species B " of 
Knowles and Das-Gupta, 1935), it is not certain that it is 
a distinct species. 

Habitat. — Gut-contents oi Hem,idactylus Haviviridis (Riippell) : 
Bengal, Calcutta. 

113. Eimeria koormse Das-Gupta. 

'fEimeria koormse, Das-Gupta, 1938, p. 155. 

Oocyst spherical, thick- walled, possessing a pseudo-micropyle 
when mature ; without oocystal residuum. Sporoblasts 
spindle-shaped ; sporocystal residuum present. 

Dimensions. — Oocysts 14 /a in diameter ; sporoblasts 10 /a by 
4-5 /i. 



Remarhs. — Full description of the form has not yet been 
published. It differs from E. legeri (Simond), described from 
the same host-species, in the form of the sporoblasts. 

Habitat. — Rectum of the Indian tortoise, Lissemys punctata 
Bonnaterre : Bengal, Calcutta. 

114. Eimeria laminata Ra5^ (Fig. 88.) 

■\Eimeria laminata, Ray, 1935, pp. 369-73, pi. x, figs. 1-11. 

Young schizonts occur in the intestinal epithelial cells ; 
liberated merozoites are active " gregarinulee," and show two 

Fig. 88. — Eimeria laminata Ray. A, a portion of the transverse section 
of the sniall intestine (a, merozoite free in the lumen ; 
b, merozoite entering a cell ; c, macroschizogony ; d, micro - 
gamete formation ; e, macrogamete ; /, microschizont ; 
g, three merozoites in a cell); B, C, macro- and micro- 
schizonts showing laminae and a siderophilous area containing 
contractile elements; D, oocyst showing sporoblasts. 
E, mature oocyst. (After Ray.) 

kinds of movements, viz., a bending movement and a screwing 
movement, in which the organism turns rapidly on its long 
axis and advances with the more pointed end forwards. This 
anterior end of the body bears on either side a hyahne blade- 
Uke structure or lamina. Schizogony is initiated at a very 
early stage and there are two types of schizonts : (a) macro - 
schizonts, releasing twenty to thirty macromerozoites, which 



develop into macrogametes, and (6) microschizonts, producing 
six to eight micromerozoites, which become microgametocytes 
and give rise to numerous uniflagellate microgametes. The 
microschizonts are distinguished by the absence of darkly 
staining granules from their cytoplasm, while the macro- 
schizonts show the granules scattered through the cyto- 
plasm from a very early stage. Sporogony is strictly intra- 
cellular. The oocyst occurs in the intestinal epithelial cell, 
is spherical, and develops four sporoblasts ; there is no oocystic 
residuum. The sporocysts are spindle-shaped, and each 
contains two sporozoites and a residuum. The mature 
oocysts are dropped into the lumen of the intestine and pass 
out in the fsecal matter. 

Dimensions. — Young schizont and merozoite 3/x by 1-25 /i 
to 12/x by 6ju. ; oocyst 8-11 ju, in diameter ; sporocyst 4*5-6 /a 
by slightly less than 3 [x. 

Remarks. — Of the 200 specimens of Bufo melanostictus 
examined only two were found to be infected, and schizogony 
and sporogony were occurring simultaneously in the same 

Habitat. — Small intestine of Bujo melanostictus Schneider : 
Bengal, Calcutta. 

115. Eimeria legeri (Simond). (Fig. 89.) 

■fCoccidium legeri, Simond, 1901 d, pp. 485-6, figs. 1-6. 
Eimeria legeri, Wenyon, 1926, p. 860 ; Reichenow, 1929, p. 950. 

Young macrogametes enter the cells of the Hver and grow 
till they reach a diameter of 16-18 ju,, when they acquire a thin 


rig. 89.- — Eimeria legeri (Simond). A, growing macrogamete ; B, en- 
cysted condition ; C, oocyst containing sporoblasts ; 
D, oocyst containing sporocysts ; E, fully developed sporo- 
cyst ; F, sporozoite. (After Simond.) 


membrane. Oocyst spherical, the cytoplasm dividing into 
four sporoblasts without leaving any residuum. The four 
sporocysts have clear contents. Each shows one smaller and 
two larger spheres ; the two larger ones represent the larger 
end of each of the contained comma-shaped sporozoites, and 
the smaller one a residual body which is granular to start with, 
but soon becomes transparent and refringent like the sporo- 

Remarks. — The parasite differs from E. kermorganti in the 
smaller dimensions of the oocysts and in the appearance of 
the sporocysts. The oocyst wall is very thin and all the 
developmental stages are passed mthin the tissues of the host. 
The sporocysts fall into the bile ducts and pass out through 
the ahmentary canal. 

Habitat. — Gall-bladder and bile-ducts of the tortoise, 
Lissemys punctata granosa (Schoepff) : India. 

116. Eimeria mitraria (Laveran & Mesnil). (Fig. 90.) 

■\Coccidium mitrarium, Laveran & Mesnil, 1902, p. 613, figs. 12-14. 
Eimeria mitraria, Wenyon, 1926, p. 859, 860, fig. 369, 3-5; 
Reichenow, 1929, p. 949. 

Schizonts produce twenty fusiform merozoites with a central 
nucleus. The microgametocytes produce a number of hair- 
like microgametes. The oocysts have a characteristic mitre- 
like form, the surface of which, unlike those of other Coccidia, 

Fig. 90.— Eimeria mitraria (Lav. & Mes.). A, oocyst with four spore- 
blasts ; B, oocyst containing four sporocysts, each with 
two sporozoites and a residue. (After Laveran and Mesnil.) 

presents four or five ornamental projections. One pole of the 
oocyst always bears a single projection, and the other pole, 
which is truncated at the base, bears three (or four) projections 
round it. The protoplasm retracts itself from the wall of the 
oocyst, becomes spherical, and divides into four sporoblasts 
without leaving any cystic residue. Each sporoblast gives 
rise to an ovoid sporocyst containing two sporozoites and a 
sporal residue. 

Dimensions. — Schizonts 10-12 jx ; merozoites 3-5 jj, in length ; 
microgametocytes 10-15 ju, in diameter ; oocysts 10-15 /x in 



Remarks. — ^AU the stages of development are extracellular, 
but are more or less intimately attached to the epithehal cells 
of the host. Laveran and Mesnil searched in vain for the 
intracellular stages, and remarked that if they exist they 
must be of a very short duration. They thought it probable 
that the organism nourishes itself at the cost of the epithehal 
cells by means of pseudopodia, and that the projections 
of the oocyst are their chitinous representations. 

Habitat. — Intestine of the tortoise, Chinemys reevesii (Gray) : 

117. Eimeria najse Ray & Das-Gupta. (Fig. 91.) 

■fEimeria najse, Kay & Das-Gupta, 1936 h, p. 345 ; 1937 d, pp. 275-7, 
pi. viii. 

Schizogony results in the formation of eight merozoites, 
belonging to two types, either spherical or oval in form. 
One type shows vacuolar cytoplasm and a distinct karyosome 

Fig. 91. — Eimeria najse Ray & Das-Gupta. A, intestinal epithelial 
cells showing (a) young male gametocyte, (6) microgamete 
formation, (c) growing macrogamete ; B, macrogamete with 
micropyle and reserve granules ; C, oocyst showing a button- 
like plug at the micropyle ; D, mature oocyst. (After Ray 
and Das -Gupta.) 

in the nucleus and later develops into female gametes. The 
other also shows hyahn cytoplasm, but the nucleus contains 
few chromatin granules besides the karysome, and later gives 
rise to male gametes. Fully formed female gametocyte shows 
a large number of reserve granules in the alveoli of the vacuolar 
cytoplasm, and a micropyle is also visible, but disappears 
when the oocyst is formed, a button-like plug being seen 
instead. Fully formed male gametocyte contains a large 
number of biflagellate male gametes. Oocysts oval, thin- 


walled. Oocystic residuum absent. Sporocysts spindle- 
shaped, with a small residue. Segmented and unsegmented 
oocysts discharged from the host. Sporulation takes place 
in 4 to 5 days. 

Dimensions. — ^Male gametocyte 20-24 /u, in diameter, female 
gametocyte 23-27 /x by 16-18 /i ; oocysts 23-27 /x by 16-18 /x ; 
sporocysts, 12-14|U, by 6-8 /x. 

Remarks. — Some of the oocysts showed sporozoites lying 
free in the oocyst, formed without the formation of sporo- 
blasts. This abnormahty has been previously described by 
Wasielewski for E. steidse and Paracoccidium prevoti. 

Habitat. — Epithelial cells of the small intestine of the cobra, 
Naja naja Linn. : Bengal, Sunderbans. 

118. Eimeria piscatori Ray & Das-Gupta. 

'\Eimeria piscatori, Ray & Das-Gupta, 1936 a, p. 345. 

Oocysts oval ; oocystic residuum present. Sporocysts 

Dimensions. — Oocysts 29-31 /x in length by 22-5-24-5;[t 
in breadth ; sporocysts 14 /x by 4-6 ju.. 

Habitat. — Rectum of the snake, Natrix piscator (Schneid.) : 
Bengal, Calcutta. 

119. Eimeria sardinas (Thelohan). (Fig. 92.) 

Coccidium sardine, Thelohan, 1890, p. 1216 ; Labbe, 1899, p. 69 ; 
Minchin, 1903, p. 340. 
■^Coccidium oxyspora, Dobell, 1919, p. 188, pi. viii, fig. 3. 

Eimeria oxyspora, Castellani& Chalmers, 1919, p. 476. 

Eimeria sp., Snijders, 1920, pp. 427-32. 

Eimeria snijdersi, Dobell, 1920, pp. 433—6. 

Eimeria oxyspora, Dobell, 1922, p. 1499, fig. 533 D ; Hegner & 
Taliaferro, 1924, pp. 289, 290, 291 ; Craig, 1926, pp. 355-7, 
fig. 64; Wenyon, 1926, pp. 851, 852, 861, figs. 350, 368; 
Knowles, 1928, p. 361, fig. 83, 1. 

Eimeria snijdersi, Wenyon, 1926, p. 855 ; Knowles, 1928, p. 361, 
fig. 83, 3. 

Eimeria sardine, Wenyon, 1926, pp. 851, 852, 861, figs. 350, 368 ; 
Thomson & Robertson, 1926 a, pp. 282-3, 2 figs. ; 1926 b, 
pp. 420-1 ; Knowles, 1928, p. 362, fig. 83, 3 ; Reichenow. 1929, 
pp. 951-2, fig. 918 A ; Kudo, 1931, p. 274, fig. 113 m. 

Oocyst spherical, having a faintly yellow transparent 
wall, composed of at least two distinct layers, and containing 
four dizoic spores and a small oocystal residue. Sporocysts 
long, sharply pointed at both ends, and possessing a tough 
endospore and deciduous epispore, the remains of which give 
the spore a frilled appearance. In each spore there are two 
sporozoites, with pointed anterior and rounded posterior 
ends, the latter containing the nucleus. 



Dimensions. — Oocyst 36-52 )u, in diameter ; sporocysts 
30-32 ;li by 7-5/Li. 

Remarks. — Dobell (1919) found a Coccidium in a young man 
who had been in South Africa, India, and Ceylon, and described 
it as a new species under the name Coccidium oxyspora. 
He could not determine whether the organism was pathogenic, 
as the patient was also infected with Entamoeba histolytica 
and Ancylostoma. He further analysed records of over seventy 
cases of coccidiosis in man. These cases all belong to Eimeria 
clupearum (Thelohan) {=E. wenyoni Dobell, 1918) or to 
Isospora belli Wenyon (=/. hominis Railhet & Lucet). 

Snijder (1920) described another case of human coccidiosis 
and, as the oocysts were definitely larger, Dobell (1920) named 

Fig. 92. — -Eimeria sardine (Thelohan). (After Dobell.) 

this parasite E. snijdersi. Broughton-Alcock and Thomson 
(1922), however, in another case of E. oxyspora infection 
found oocysts quite as large as those of E. snijdersi, and con- 
sidered the two species to be identical. Brug (1922 a) 
suggested that E. snijdersi was of animal origin and ingested 
with food. Thomson and Robertson (1926 a) made a careful 
study of the Coccidia of fish and came to the conclusion that 
the oocysts of E. sardine (Thelohan, 1890), parasitic in the 
" soft roe " (testis) of herrings, sprats and mackerel, are 
identical with the oocysts of E. oxyspora and E. snijdersi. 
They further (1926 b) proved their contention by giving 
a man a strong sahne aperient and afterwards making him 
eat a large quantity of soft roe of herring. Next morning he 
passed a large number of oocysts of E. sardinm which were 



identical with those found in cases of human coccidiosis. 
Thus E. oxyspora and E. snijdersi are synonymous with 
E. sardine. 

In India Setna and Bana (1935) have studied the coccidial 
infection in a number of fish, and described the oocysts and 
sporocysts from ten different species, but the oocysts in no 
case correspond with those of E. dujjearum or E. sardinse. 

Habitat. — ^AHmentary canal of man who had visited South 
Africa, India, and Ceylon. 

120. Eimeria smithi Yakimoff & Galouzo. (Fig. 93.) 

Coccidium oviforme (part), Guillebeau, 1893, p. 81. 

Eimeria zurni (part), Smith & Graybill, 1918, p. 89 ; Wenvon, 

1926, pp. 842-3, fig. 362. 
fEirneria zurni (part). Cooper, 1926 a, p. 290 ; 1926 b, p. 291. 
fBovine Coccidia (part). Cooper, 1927, pp. 92-7, pi. x. 
Eimeria smithi, Yakimoff & Galouzo, 1927, pp. 185-200, figs. 1-7 ; 
Reiehenow, 1929, p. 945 ; Rastegaieff, 1930, pp. 390-1, fig. 2. 
fBovine Coccidia, Sen, 1932, p. 34. 
^Eimeria smithi. Ware, 1936, p. 35. 

Oocysts are ovoid, with one pole pointed, have a thin wall, 
and are provided with a micropyle. They are brownish in 
colour, and are distinctly larger in size than those of E. zilrni. 
Mature cysts do not show any oocystic residue. Sporocysts 
pear-shaped, with a sporo cystic residue. 

Fig. 93. 

Fig. 95. 

Fig. 94. 

Fig. 93. — Eimeria smithi Yakimoff & Galouzo. (After Yakimoff and 

Fig. 94. — Eimeria wassilewskyi Rastegaieff. (After Rastegaieff.) 
Fig, 95. — Eimeria zurni (Rivolta). (After Yakimoff and Galouzo.) 

Dimensions. — Oocysts 25-35^ in length; sporocysts 10-8- 
14-4 jLt by 7-9 /u. ; sprozoites 3-6-5-8(U, by S-hfi. 

EemarJcs. —Zscho^e (1892), Hess (1892), and Guillebeau 
(1893) were the first to describe coccidiosis of cattle as a 
distinct disease. Guillebeau noted that in certain years the 
disease became epidemic and caused considerable mortahty. 
Multiplication occurred in the epithelial cells of both the small 


and large intestine. The oocysts described by him possessed 
a distinct micropyle. ZubHn (1908) studied the development 
of the oocyst outside the body and also noted that although the 
majority of oocysts were 12-15 )u. in diameter, larger forms, 
which measured 30-35 ju, in length by 20 ft in breadth, also 
occurred. Theobald Smith and Graybill (1918), who investi- 
gated coccidial dysentry of calves in America, also encountered 
oocysts of two types. The second t3^e, which evidently 
corresponds with E. smithi, were said to be brownish in colour, 
possessed a thick wall, and measured 25-8-41-8/x in length by 
16-4-24-6/x in breadth. They did not contain any oocystic 
residue, but sporocystic residue was present. 

Wenyon (1926) considered it possible that there were 
two species of Eimeria found in cattle, but was led to identify 
and figure the larger, oval type as E. zilrni. 

Cooper (1924) apparently was the first to record bovine 
coccidiosis in India. He (1926 a) noted the extreme variation 
in shape and size of the oocysts, but considered these variations 
to be connected with the rate of multiplication. According 
to him, in normal "carrier" infection all oocysts are weU 
formed, with a strong, thick capsule and with a relatively 
large amount of dense and granular protoplasm, but very large 
and extremely small forms are encountered, and display 
considerable range of variation in shape. At the height of 
multipHcation oocysts are small, and they show an almost 
uniformly ovoid to a nearly round shape, with distinctly 
thinner capsules. When clinical symptoms occur still smaller 
oocysts appear, usually in smaU numbers ; these have an 
extremely thin capsule and much reduced amount of almost 
transparent protoplasm. In discussing the pathogenicity of 
bovine Coccidia, Cooper (1926 b) maintained that a latent or 
" carrier " type of infection is of almost universal occurrence 
in cattle in India, and that, although usually imiocuous, 
the parasites are capable, under certain conditions, of over- 
coming the animal's natural defences and then set up a cHnical 
disease identical with what is described in other countries as 
" red dysentery." He also observed that clinical coccidiosis 
also occurs as a sequel to rinderpest, even a mild attack of 
which may result in an appreciable resuscitation of Coccidia. 
Later (1927) he again reviewed bovine coccidiosis, but the 
possibihty of the existence of two species did not occur to him. 
YakimofiF and Galouzo (1927) surveyed the previous htera- 
ture on bovine Coccidia and gave a table noting the measure- 
ments of oocysts and other character as given by previous 
workers, and came to the conclusion that the name E. ^iirni 
should be restricted to the forms with smaller and spherical 
oocysts, while larger and oval oocysts should be referred to 
a new species, E. smithi. 



Sen (1932) recorded a case of coccidiosis in a buffalo calf 
in the Coorg District, and stated that the parasites comprised 
two species, one a larger form, of yeUow colour, provided with 
a micropyle and the other a small colourless form without 
a micropyle. I think the former can be identified as E. smithi 
and the latter as E. zilrni. Ware (1936) has for the first time 
noted the occurrence in India of ^. smithi as such. 

Habitat. — AHmentary canal of Bos indicus Linn. (?) : 
United PnoAncNCES, Muktesar ; ahmentary canal of Bos 
bubalus Linn. : South India, Coorg. 

121. Eimeria southwelli Halawani. (Fig. 96.) 

"fEimeria southwelli, Halawani, 1930 a, pp. 1-3, fig. 1; 1930 6, 
p. 326. 

Oocysts polymorphic. Immature oocysts generally pea- 
shaped. Mature oocysts cylindrical or sausage-shaped ; 

Fig. 96. — Eimeria southwelli Halawani. A, an immature pear-shaped 
oocyst containing a large spherical zygote ; B, an imma- 
ture oocyst retaining its hind-bulb to a slight degree ; 
C, oocyst containing four sporoblasts ; D, fully matm-e 
oocyst containing oval sporocysts. (After Halawani.) 

cyst- wall colourless and transparent. A residual mass is 
usually present in the immature cyst, but often disappears as 
the cyst matures. Sporoblasts four, arranged lengthwise, 
end to end, sometimes in pairs or in a chain. Sporocysts oval. 

Dimensions. — Oocysts 25-50 )u, in length, average dSfx, 
average width 12 ^u. ; sporocysts 10-12 /a in length. 

Remarks. — It is presumed by Halawani that the matme 
oocysts from the parent pass, via the cloaca, up the oviduct 
to the uterus, and by hberatton of their sporozoites infect 
the intra-uterine embryos. 

Habitat. — Spiral valve of the intestine of the embryo of 
a shark, Mtobatis narinari Agassiz, from the Indian Ocean : 
Ceylon, Colombo. 

SPOE. o 



122, Eimeria wassilewskyi RastegaieflF. (Fig. 94.) 

■fEimeria wassilewskyi, Rastegaieff, 1930, pp. 391-2, fig. 3. 

Oocysts egg-shaped, flattened at one end, with a distinct 

Dimensions. — Oocysts 18 /a by 14-4 ju, ; micropyle 4-5 /a. 

Habitat. — ^Alimentary canal of Axis axis Erxl. {=Gerviis axis), 
from India, in the Zoological Gardens at Leningrad. 

123, Eimeria yakimovi Rastegaieff. (Eig. 97.) 

■fEimeria yakimovi, Rastegaieff, 1930, p. 389, fig. 1, 

Oocysts oval, with distinct micropyle and containing 
four sporocysts. Sporocysts oval, each containing two sporo- 

Fig. 97.- 

-Eimeria yakimovi Rastegaieff. 
(After Rastegaieff.) 

zoites, which are pyriform, and lie in a tete-beche manner. 
No cystal or sporal residue present. 

Dimensions. — Oocysts 32-4-41-4ju, by 21-6-28-8jLt ; sporo- 
cysts 13-19)Lc by 3-5-4ju. ; sporozoites lI-7-13'6ju, by 4-5/x. 

Habitat. — Alimentary canal of the nilgai, Boselaphus 
tragocamelus (Pall.), from India, in the Zoological Gardens at 

124. Eimeria ziirni (Rivolta). (Fig. 95.) 

Coccidium perforans (part), Ziirn & Proger, 1877, p. 113. 
Cytospermum, ziirni, Rivolta, 1878. 
Coccidium ziirni, RailHet & Lucet, 1891, p. 247, 
Coccidium perforans var., Labbe, 1899, p. 67. 

Eimeria ziirni, Jowett, 1911, p. 207 ; Smith & Graybill, 1918, 
p. 89 ; Hegner & Taliaferro, 1924, p. 287 ; Wenyon, 1926, 
pp. 842-3. 
IBovine Coccidia, Cooper, 1924, p. 48. 
•fEimeria ziirnii, Cooper, 1926 a, p. 290. 
Eimeria ziirni, Yakimoff & Galouzo, 1927, pp. 185-200, figs. 8-15; 
Reichenow, 1929, p. 945 ; Rastegaieff, 1930, p. 399. 
tSmaller form, Sen, 1932, p. 34. 


Oocysts colourless or faintly greenish, almost or quite 
spherical, without a micropyle, and smaller than those of 
E. smiihi. Mature cysts contain no oocystal residue, and the 
sporoblasts, when first formed, are spherical. They soon 
become ovoid and secrete a sporocyst which has a thick cap 
at one pole. The sporocysts are completely filled by sporo- 
zoites and there is no sporocystal residue. 

Dimensions. — Oocysts 12-25 jit in diameter ; sporocysts 9-9- 
lljuby 5-3-5-7)Li. 

Remarks. — Zublin (1908) was the first to differentiate the 
larger oocysts found more rarely (now identified as E. smithi) 
from the smaller ones which are more commonly found, and 
which measure 12-25 in diameter. Jowett (1911) gave the 
measurements as 14-4-27-2 yu, by 12-8-20-8ju,. Theobald Smith 
and Graybill (1918) also described two tjrpes of oocysts, and 
gave the measurements of the smaller ones as 13- 1-28-7 yn by 
12- 3-20-5 ja. 

Habitat. — AHmentary canal of Bos indicus Linn. (?) : 
United Provinces, Muktesar ; aHmentary canal of Bos 
huhalus Linn. : South India, Coorg. 

125. Eimeria sp. 

■\Eimeria sp., Cooper & Gulati, 1926, pp. 191-2. 

Oocysts of an unidentified species of Eimeria were found 
in the faeces of a cow at Tocklai and a heifer at ShUlong, in 


126. Eimeria sp. 

■fEimeria sp., Setna & Bana, 1935 a, p. 167. 

Oocysts spherical ; residual body absent. Sporocysts 
ovoid, with a small rounded knob-Uke thickening at one end ; 
sporocystal residue absent. 

Dimensions. — Oocysts 10-6 /x in diameter ; sporocysts 5-3 ft 
by 3 (J,. 

Habitat. — Intestine of the fish Trichiurus savala Guv. & Val. : 
Bombay, Bombay. 

127. Eimeria sp. 

■\Eiineria sp., Setna & Bana, 1935 a, p. 167. 
Oocysts spherical ; residual body occasionally present. 
Sporocysts small ovoid, with one pole sKghtly narrower 
than the other ; sporocystal residue absent. 

Dimensions. — Oocysts 7-6 /x in diameter; sporocysts 3-4 /i 
by 2-1 /x. 

Habitat. — Intestine of the fish Batrachus grunniens (Bl. «& 
Schn.) : Bombay, Bombay. 



128. Eimeria sp. 

■\Eimeria sp., Setna & Bana, 1935a, p. 167. 

Oocysts spherical ; residual body absent. Sporocysts 
ovoid ; the sporocystal residue in the form of two darkly 
stauiing globules. 

Dimensions. — Oocysts 10 /a in diameter: sporocysts 5-5 /x, by 
3-1 /x. 

Habitat. — Intestine of the Q.8h.Epinephelustauvina (Forskal) : 
Bombay, Bombay. 

129. Eimeria sp. 

■f Eimeria sp., Setna & Bana, 1935 a, p. 167. 

Oocysts spherical ; residual body absent. Sporocysts 
broadly ovoid ; sporocystal residue absent. 

Dimensions. — Oocysts 12jli in diameter ; sporocysts 7-6 /x, by 
6-1 /x. 

Habitat. — Intestine of the fish Etigraulis mystax (Bl. & 
Schn.) : Bombay, Bombay. 

130. Eimeria sp. 

■fEimeria sp., Setna & Bana, 1935 a, p. 167. 

Oocysts spherical, residual body absent. 
Dimensions. — Oocysts 10-3 /x in diameter. 
Habitat. — Intestine of the fish Otolithus ruber (Bl. & Schn.) : 
Bombay, Bombay. 

131. Eimeria sp. 

■fEimeria sp., Setna & Bana, 1935 a, p. 167. 

Oocysts almost spherical ; residual body absent. Sporo- 
cysts ovoid, with one end more pointed than the other ; 
sporocystal residue absent. 

Dimensions. — Oocysts 8-4 /n in diameter ; sporocysts 4/x by 
2-5 fi. 

Habitat. — ^Intestine of the fish Sillago sihama (Forskal) : 
Bombay, Bombay. 

132. Eimeria sp. 

■^Eimeria sp., Setna & Bana, 1935 a, p. 167. 

Oocysts spherical ; residual body absent. Sporocysts 
ovoid, with frequently a knob at one end ; sporocystal 
residue absent. 

Dimensions. — Oocyst 15-18 ju, in diameter ; sporocysts 8-7 ju. 
by 5-3 ju.; 

Habitat. — Intestine of the fish Coilia dussumieri (Cuv. & 
Val.) : Bombay, Bombay. 


133. Eimeria sp. 

^Eimeria sp., Setna & Bana, 1935 a, p. 167. 

Habitat. — Intestine of the fish Plotossus canius (Ham.-Buch.) : 
Bombay, Bombay. 

134. Eimeria sp. 

■\Ei->neria sp., Setna & Bana, 1935 a, p. 167. 

Oocysts spherical ; residual body absent. Sporocysts 
ovoid ; sporocystal residue absent. 

Dimensions. — Oocysts Il/x in diameter ; sporocysts 4-2 /x, by 

Habitat. — Intestine of the fish Epinephelus diacanthus 
(Cuv. & Val.) : Bombay, Bombay. 

135. Eimeria (?) sp. 

Certain Coccids, Ross, 1898, p. 173 ; 1906, pp. 102, 104. 

Well-defined oval organisms, 8ju, by 4/i in size, were found 
lying by the side of the nucleus within the stomach-cells of 
a few individuals of Culex sp. Each possessed a vacuole (?) 
surrounded by a faint granulation. According to Ross they 
probably belonged to the Coccidiidse, but were never seen again 
by him. 

Habitat. — Stomach-cells of Culex sp, : Madras, Ootacamund. 

Genus WENYONELLA Hoare, 1933. 

Wenyonella, Hoare, 1933, pp. 369-74; Ray, 1935, pp. 112-13; 
1937, pp. 117-20. 

Oocysts develop four sporocysts, each containing four 

Eemarks. — Hoare (1933) founded this genus for a species 
from the small intestine of a snake in Uganda. Ray (1935, 
1937) has described a new species from a squirrel. 

136. Wenyonella hoarei Ray. (Fig. 98.) 

"^Wenyonella hoarei, 'Raj, 1935, pp. 112-13, 6 figs. ; 1937, pp. 117- 
20, pi. vii. 

Schizonts more irregular in shape than the sexual forms, 
and lacking a clear outhne. Merozoites in groups of six 
to eight, escaping from a host-cell into the lum.en of the 
intestine. Two types of merozoites : one with hyahne cyto- 
plasm and a dark- staining granule in the neighbourhood of 
the nucleus ; the other slightly longer, more opaque when 
stained, and showing numerous dark granules scattered 



irregularly. On entering new epithelial cells the merozoites 
assume a round form and become gametocytes, which are 
dififerentiated into microgametocytes and macrogametocytes 
respectively ; the former give rise to a large number of 
biflagellate microgametes round a small cytoplasmic residuum ; 
they then break off from the residuum and swim actively, 
congregating at one end of the female gamete. The latter 
increase in size and become macrogametes. 

Oocysts perfectly spherical ; tetrasporocystid. Sporocysts 
show a characteristic lenticular knob at one pole ; tetrazoic. 
Sporozoites more or less regularly arranged, with broad ends 
of two sporozoites at each pole. 

Dimensions. — Merozoites 6/i by 2/x or 8/x by 2[x; young 
gametocytes 6-8 yu, ; oocyst 14-18-5 ju. in diameter ; sporocysts 
10/Li by S/i. 

Remarks. — In a fresh preparation merozoites were seen 
actively swimming about with the pointed end directed 

Fig. 98. — Wenyonella hoarei Ray. A, oocyst with developing sporo- 
blasts ; B, oocyst with distinct sporoblasts ; C, mature 
oocyst showing sporocysts each containing foiir sporozoites. 
(After Ray.) 

forward. Female gametocyte showed a micropyle which 
did not persist in the oocyst. Development of the male 
gametes was also observed. Advanced male gametes were 
seen adhering round a central mass of cytoplasm with their 
actively moving tail-ends directed away from it. Within 
an hour and a half these gametes broke off and congregated 
round the micropylar end of the female gamete. 

On keeping the oocysts in 1 per cent, chromic acid these 
showed the first sign of development after forty-eight hours, 
when the sporoblasts were seen to be budding out. They 
separated from each other on the fourth day, were invested 
with sporocysts on the fifth day, and the sporozoites were 
well differentiated on the seventh. 

Habitat. — Gut-contents of a squirrel, Sciurus sp. : Bengal, 


Subfamily BARROUSSIIN^ (Wenyon, 1926). 

Oocysts contain many sporocysts, each containing one or 
four sporozoites. 

Genus PYTHONELLA Ray & Das-Gupta, 1937. 
Pythonella, Ray & Das-Gupta, 1937, p. 2. 

Oocyst develops sixteen sporocysts, each containing four 

137, Pythonella bengalensis Ray & Das-Gupta. 

fPythonella bengalensis, Ray & Das-Gupta, 1937, p. 2, 

Oocysts spherical. Segmenting oocysts found in the 
epitheHal cells of the intestine and in the fseces. Oocyst 
develops eight primary sporoblasts, and after some time these 
divide to form sixteen, and are converted into as many sporo- 
cysts. Each sporocyst contains four sporozoites and a central 
residuum . Male gametes are formed as in the family Eimeriidse . 
Female gametes often seen lying in the submucosa. 

Dimensions. — Oocysts 25-30 fi in diameter ; sporocysts 8-10 /i 
by 6-7^. 

Remarks. — Full description of this species has not yet been 
pubhshed. Heccaidecasporocystid and tetrazoic condition 
marks it out from all known Coccidia. Segmented or unseg- 
mented oocysts were seen in the faecal matter and matured 
within seven to ten days when kept in 1 per cent, chromic acid. 

Habitat. — Intestine of Python sp. : Bengal, Calcutta. 

Subfamily AGGREGATING Reichenow, 1929 
(emend. Hoare, 1933). 

Schizogony in one type of host and sporogony in another. 
Oocysts contain many sporocysts, each sporocyst giving 
rise to from two to many sporozoites. 

Remarks. — Labbe (1899) founded the family Aggregatidse 
to include the genus Aggregata. As originally defined by 
Frenzel, Aggregata was a genus of Gregarines, characterized 
by sporozoites being formed directly in the cyst round a number 
of residual masses. The researches of Dobell (1914, 1925) 
and Pixell-Goodrich (1924) showed that the Aggregatidse may 
be safely regarded as Coccidia, which have their schizogony 
in one host (usually a crab) and the sporogony in another 
(usually a Cephalopod). It is now known that the bodies 
produced in the cyst are merozoites, and not sporozoites 
as was formerly supposed. 


Reichenow (1929) amended the family and included in it 
a number of other genera, such as Pseudoklossia Leger & 
Duboscq, Merocystis Dakin, Myriospora Lermantoflf, Caryo- 
tropha Siedlecki, and Angeiocystis Brasil, all parasites of marine 
worms, molluscs and crustaceans, in which the oocyst develops 
many sporocysts (except in Angeiocystis Brasil, in which only 
four are developed) containing two or more (up to thirty) 
sporozoites. In some of these genera schizogony is not known, 
and therefore is presumed to take place in some other host. 
Hoare (1933), in accordance with his scheme of classification, 
has transferred Angeiocystis to the subfamily Eimeriinse, 
and has retained the others in his subfamily Aggregatinse. 
Mackinnon and Ray (1937) have shown that the sporozoan 
parasite hitherto known as Monocystis ihalassemse Lankester 
is a coccidian, and have placed it in a new genus, Ovivora, 
belonging to this subfamily. 

Genus AGGREGATA Frenzel, 1885. 

" Sac a Psorospermies," Van Beneden, 1854, p. 9. 

Monocystis sp., Lieberkiihn, 1855, p. 9, pi. viii, figs. 9, 12. 

Benedinia, Aime Schneider, 1875 a, pp. xl-xlv. 

Klossia, Aime Schneider, 1883, pp. 78-104, pis. viii, ix. 

Aggregata, Frenzel, 1885, p. 560. 

Gregarina, Frenzel, 1885, pp. 572, 576, 578. 

Benedinia, Labbe, 1895, p. 381 ; 1899, pp. 54-5. 

Klossia, Labbe, 1896, p. 535, pi. xii, fig. 20 ; pis. xv, xvi, xviii, 
figs. 1-12; 1899, p. 54. 

Aggregata, Labbe, 1899, p. 6. 

Leger ia, Blanchard, 1900, p. 159. 

Eucoccidium, Luhe, 1902, pp. 771-3. 

Legerina, Jacquemet, 1903, pp. 193-4. 

Aggregata, Leger & Duboscq, 1906, pp. 1001-3 ; 1908, pp. 44-108 ; 
Moroff, 1906 a, pp. 652-4 ; 1908, pp. 1-224 ; Pixell-Goodrich, 
1914, pp. 159-74; Dobell, 1914, pp. 1-7; 1925, pp. 1-136; 
Wenyon, 1926, pp. 870-5, figs. 376-8; Knowles, 1928, pp. 366-8; 
Reichenow, 1929, pp. 928-31, figs. 894-7 ; Kudo, 1931, p. 270, 
fig. Ill ; Calkins, 1933, p. 566; Reichenow, 1935, p. 373. 

Oocyst contains many sporocysts, each developing three 
sporozoites. Schizogony takes place in a crab, and sporogony 
in a cephalopod host. 

Remarks. — The life-cycle of ^. eberthi Labbe has been studied 
by a number of workers. The nucleus of the zygote divides 
repeatedly, and numerous sporoblasts, and finally sporocysts, 
are developed in the oocyst in the body of a cuttlefish. Each 
sporocyst contains three sporozoites and a residual mass. 
A crab is infected by eating the infected material passed in 
the dejecta of a cuttlefish. The sporozoites are liberated 
in the intestine of the crab and, passing through the lining 
cells, grow into schizonts, which form cysts that bulge into 
the body-cavity, and by schizogony produce innumerable 



Fig. 99. — Life-cycle of Aggregata eherthi Labbe. The stages above 
the dotted line occur in the cuttlefish, those below in the 
crab. R, merozoite swallowed by the cuttlefish ; A, un- 
differentiated parasite in submucous tissue ; B, C, D, growth 
into niicrogametoe3rte and production of microgametes ; 
E, F, G, growth into macrogamecyte and fertilization ; 
H, zygote ; /, first nuclear division in zygote ; J, K, nuclear 
multiplication in zygote and production of sporoblasts ; 
L, sporocyst containing three sporozoites and small residual 
body ; M, escape of sporozoites in intestine of crab ; N-Q, 
growth of schizont and production of merozoites in sub- 
epithelial connective tissue. (From Wenyon, after DobeU.) 


merozoites. When the crab is eaten by a cuttlefish, the 
merozoites penetrate the gut-wall and develop into micro- 
and macrogametocytes, and further into gametes. Anisogamy 
results in zygote formation, and the oocysts are passed out 
and ingested by another crab. 

138. Aggregata sp. 

•fAggregata sp., Setna & Bhatia, 1934, pp. 42-3, fig. 23. 

Schizonts spherical or oval, and are enclosed in a cyst. 
Each cyst has six or seven pear-shaped apertures through 
which merozoites escape. The merozoites are curved spindle- 
shaped bodies, with a broad central portion and pointed ends ; 
the nucleus consists of deeply staining granules and occupies 
one end. The merozoites move with sudden jerky or springy 
movements, and the nuclear end is usually anterior in these 

Dimensions. — Schizonts 193-246 /u. in diameter; merozoites 
ll-16ju. in length. 

Habitat. — Intestine of the prawn, Parapeneopsis sculptilis 
(Heller) : Bombay, Bombay. 

Incert^ sedis. 
Genus TOXOPLASMA Nicolle & Manceaux, 1909. 

Toxoplasma, Nicolle & Manceaux, 1909 a, pp. 369-72 ; 1909 6, 
pp. 97-103; Splendore, 1909, pp. 462-5; Carini, 1909, pp. 465-9 ; 
1911 a, p. 518 ; Ugo Mello, 1910, pp. 359-63; Yakimoff & Kohl- 
Yakimoff, 1911a, pp. 617-18; 1912, pp. 1-14; Bourret, 1911, 
p. 373 ; Sangiorgi, 1913, p. 323 ; 1914, p. 83 ; MaruUaz, 1913, 
pp. 323-6 ; Laveran& MaruUaz, 1914, pp. 21-5 ; Castellani, 1914, 
p. 113 ; Castellani & Chalmers, 1919, pp. 488-90 ; NoUer, 1920, 
pp. 907-18; Hegner & Taliaferro, 1924, pp. 293, 300-2; Brug, 
den Heyer, & Haga, 1925, pp. 232-8. 

Babesia (Toxoplasma), Calkins, 1926, p. 445. 

Toxoplasma, Wenyon, 1926, pp. 1041-50; Knowles, 1928, pp. 463-5; 
Reichenow, 1929, p. 965 ; NoUer, 1931, pp. 804-7. 

Organism small, elongated, shghtly curved, with a central 
nucleus. Found in the host-cells either singly or in groups 
resulting from repeated binary fission. When occurring 
singly they often he against the nucleus of the host-cell and 
indent it, thus bearing some resemblance to the leucocytic 
Hsemogregarines. Parasites of body fluids, leucocytes, and 
cells of the spleen, liver, kidneys, lungs, etc., of various 

Species have been described from man, monkeys, dogs, 
gondis, rabbits, rats, guinea-pigs, moles, birds, and snakes. 

Remarks. — It is not possible to determine the correct 
position of Toxoplasma in the scheme of classification till more 


is known about the life-history. Calkins (1926) provisionally- 
regarded it as a subgenus of Babesia, while Wenyon (1926) 
placed it among parasites of doubtful nature. Reichenow 
(1929) placed the genus as an addendum to the family Eimeridse, 
as, according to NoUer, certain stages found in the wall of 
the ahmentary canal of birds may be interpreted as schizonts, 
microgametocytes, and macrogametes of the type found in 
that family. Noller (1931) thinks that they should most 
probably be placed in the neighbourhood of the Coccidia 
occurring in the blood. Kudo (1931) and Calkins (1933) 
have altogether excluded them from consideration. 

Key to Indian Species. 

1 (8). Schizogony known to occur 2. 

2 (7). Schizonts not diHerentiated 3. 

3. Form circular or pointed, 3-7 /x ; schizo- 

gony in the Uver-eells, more rarely in 

peripheral blood or bone-marrow ; 

sometimes binary fission. In leuco- [Kohl-Yak., p. 204. 

cytes of pigeon T. colwmbee Yak. & 

4. Smaller forms ring-like ; schizonts 

showing signs of binary fission. In [p. 203. 

Butastur T. butasturis de Mello, 

5. Form, crescentic, pointed at both ends, 

reproduction by binary fission or 

schizogony. In smears from internal [p. 205. 

organs of rabbit T. cuniculi Splendore, 

6. Form semilunar or oval. In cells of [p. 204. 

internal organs of dog T. canis Ugo Mello, 

7 (2). Schizonts and merozoites of two kinds : 

form ovoid, fusiform or falciform. 

In leucocytes or endothelial cells of [p. 206. 

lung of coot T. fulicse de Mello, 

8 (1). Schizogony not known, even animal 

natm-e doubtful. Form round, ovoid 

or pyriform. In blood and spleen of [lani, p. 207. 

man T. {!) pyrogenes Castel- 

139. Toxoplasma butasturis de Mello. 

■f Toxoplasma butasturis, de Mello, 1937 a, p. 111. 

Small parasites resemble the ring-forms of Plasmodids, 
larger schizonts with the nucleus better formed and showing 
evidence of binary fission. Often more than one parasite 
in the same host-cell. 

Bemarks. — The organism differs from T. fulicsa in that 
the cytoplasm is stained uniformly pale rose and does not 
show two types of coloration suggestive of sexual differentia- 
tion. It resembles T. columhse Yakimoff & Kohl-Yakimoff, 
of the pigeon. 

Habitat. — Leucocytes of Butastur teesa (FrankHn) : Portu- 
guese India. 



140. Toxoplasma canis Ugo Mello. 

Toxoplasma canis, Ugo Mello, 1910, pp. 359-63 : Minchin, 1912, 

p. 389; Carini & Maciel. 1913, pp. 681-3; Castellani & Chalmers, 

1919, p. 489; Boez, 1921, pp. 479-82; Wenyon, 1926, p. 1047. 

■f Toxoplasma canis, Donovan (first recorded in Wenyon, 1926, 

p. 1357). 

In fresh preparations the organisms are transparent, 
colourless, immobile, semilunar or oval in form, rarely rounded 
or pyriform. Stained with Giemsa each organism is a clear 
blue homogeneous mass, surrounded by a transparent and thin 
envelope. It contains one or (rarely) two chromatic masses 
representing the nucleus, which are stained reddish- violet ; 
generally rounded, but sometimes irregular, oval or linear in 
form. The organism occurs singly or in groups of any number 
up to sixty in endothelial cells, leucocytes, or cells of the spleen 
or kidney. 

Habitat. — In the endotheHal cells or cells of the spleen or 
kidney of the dog : India (locality not cited). 

141. Toxoplasma eolumbsB Yakimoff & Kohl-Yakimoff. 

(Fig. 100.) 

Toxoplasma columbse, Yakimoff & Kohl-Yakimofi, 1912, p. 198. 
"fHaemogregarina francae, de Mello, 1915, pp. 93-4, pi. ii, figs. 1-10. 
fLeucocytogregarina francee, de Mello, de Sa, de Sousa, Dias, & 
Noronha, 1917, p. 13. 

Toxoplasma francee, NoUer, 1920, p. 914. 

Toxoplasma columhse, Noller, 1920, p. 914. 

Toxoplasma francse, Wenyon, 1926, pp. 1043, 1047. 

Toxoplasma columbse, de Mello, 1935 a, p. 706 ; 1937 a, p. 111. 

Parasite has a circular form when at rest, but shght move- 
ments in the interior of the white cell change the form to a 

Fig. 100. — Toxoplasma columhse Yak. & Kohl -Yak. 
A, young form ; B, C, schizogony. (After de Mello.) 

pointed one, with the nucleus in the rounded portion. Nucleus 
in the fresh state strongly refringent, surrounded by such 
clear protoplasm that it has the appearance of a vacuole. 



Stained by Giemsa the nucleus is coloured a brilliant red ; 
the cytoplasm clear and faintly blue. Small or large forms, 
3-7 fi in size. Schizogony in the hepatic cells, more rarely 
in the peripheral blood and bone-marrow. Sometimes 
binary division takes place, when rudimentary karyokinetic 
figures can be demonstrated. Sporogony not known. 

Habitat. — Mononuclear leucocytes of the pigeon : Portu- 
guese India, Mapu9a (Bardez). 

142. Toxoplasma cuniculi Splendore. (Fig. 101.) 

Toxoplasma cuniculi, Splendore, 1909 a, p. 109 ; 1909 b, 
pp. 462-5: Carini, 1909, pp. 465-9, figs. 1-14; 1910, p. 167; 
1911a, pp. 518-19; Bourret, 1911, p. 373; Castellani & 
Chalmers, 1919, pp. 489, 490; Brug, den Heyer, & Haga, 1925, 
pp. 232-8, pis. i-vi ; Wenyon, 1926, p. 1047 ; Reichenow, 1929 
p. 965 ; ISToller, 1931, p. 805. 
'^Toxoplasma ctXniculi, Krishnan & Lai, 1933, pp. 1049-50, pi. xlv. 

Intracellular in the endothelial cells, or leucocytes, or in the 
body fluids. Crescentic in shape, pointed at both ends ; 
but one end is frequently more pointed than the other. Nucleus 
central and has a definite karyosome. Occurs singly or in 
groups. Reproduction by schizogony has also been described. 

Remarks. — Krishnan and Chiranji Lai (1933) found this 
infection in two out of twelve rabbits that had received a course 

Fig. 101. — Toxoplastna cuniculi Splendore. A, a large cluster of parasites. 
B, a large mononuclear cell from spleen showing a dividing 
form and a group of parasites. (After Krishnan and Lai.) 

of fifteen injections of Indian ink and colloidal iron, followed 
by three intravenous injections of a culture of Leishmania 
donovani. Eight weeks after the last injection the two rabbits 
began to look sickly and lose weight. They suffered from 
diarrhoea and died. Smears from spleen, Hver, bone-marrow, 
and heart-blood showed large numbers of intracellular and 



extracellular forms of this parasite. The parasite is mor- 
phologically indistinguishable from T. gondi. Splendore 
(1909) was able to infect rats, guinea-pigs, rabbits, and 
frogs, while Carini (1909) infected pigeons with this parasite. 

Habitat. — Smears from Hver, spleen, bone-marrow, and heart- 
blood of rabbit, Lepus sp. : Bengal, Calcutta. 

143. Toxoplasma fulicse de Mello. (Fig. 102.) 

■fToxoplasma fulicee, de Mello, 1935 a, pp. 708-9, pi. ii ; 1937 a, 
p. 111. 

Free, or included in endothelial cells or mononuclear leuco- 
C3rtes. Form ovoid, fusiform or falciform in young intra- 
cellular stages ; large ovoid, occupying most of the host-cell 
when full grown, but not invading its nucleus. Schizonts 
show sexual differentiation. In one type the cytoplasm is 

Fig. 102. — Toxoplasma fulicse de Mello. A, jB, female schizonts; G,D, 
division stages ; E, F, male schizonts ; O-I, division 
stages. (After de Mello.) 


stained dark blue and the nucleus is a compact chromatic 
dot, sometimes vacuolated, and surrounded by a white 
halo : in the other type the cytoplasm is stained light violet, 
rather rosy, and the nucleus is compact or vacuolated, some- 
times constituted by an irregular spirematic thread, never 
showing the vacuolar space round it. The former are inter- 
preted as female and the latter as male schizonts. Multiplica- 
tion takes place by repeated binary fission. Merozoites pro- 
duced also show the same cytoplasmic differentiation as the 
parent schizonts. 

Remarks. — The parasite was found only in smears from the 
lung, and was not found in the peripheral blood or in smears 
from other organs. 

Habitat. — Lung-smears of Fulica atra Linn. : Portuguese 
India, lakes of Carambolin and Taleigao. 

144. Toxoplasma spi 

fA parasite in the sparrow, J. R. Adie, 1908, p. 176. 
Toxoplasma avium, Castellani & Chalmers, 1918, p. 489. 
Toxoplasma sp., Wenyon, 1926, pp. 1042, 1048 ; Knowles, 1928, 
p. 465. 

Remarks. — Wenyon (1926) regards the form as undoubtedly 
a species of Toxoplasma. 

Habitat. — Blood of the sparrow, Passer sp. : Punjab. 

145. Toxoplasma sp. 

■\Toxoplasm,a sp., Plimmer, 1916 a, p. 295. 

^Toxoplasma sp., Wenyon, 1926, pp. 1043, 1048 ; de Mello, 1937 a, 
p. 111. 

Habitat. — Blobd and exudation from the lungs of the chat, 
Saxicola caprata Linn., from India, in the Zoological Gardens, 

Doubtful Species. 

146. Toxoplasma (?) pyrogenes Castellani, 1914. (Fig. 103.) 

■\Toxoplasma pyrogenes, Castellani, 1914 6, pp. 113-14, 2 pis. 
Some tmknown protozoon, Plate, 1914. 
Contaminating organisms of a vegetable nature, Wenyon, 1923, 

pp. 527-50, 855, 1 pi. ; 1926, p. 1048, fig. 433, A-C. 
Toxoplasma pyrogenes, Knowles, 1928, p. 465 ; Reichenow, 1929, 

p. 965 

Intracellular, round, ovoid or pyriform in shape, and about 
6)u, in diameter. A second form, 12/x in diameter, and con- 
taining several chromatin granules, has been described as 
a schizont. 

Remarks. — ^Wenyon (1923) discussed the probable nature 
of this form and came to the conclusion that what had been 


described as T. pyrogenes was only a contaminating organism 
of a vegetable nature. Knowles (1928) and Reichenow 
(1929) support this view. Knowles has also been struck 

fe Ik 

D E 

Fig. 103. — Toxoplasma (?) pyrogenes Castellani. A-D, small forras; 
E, large form described as schizont. (From Wenyon, after 
Castellani. ) 

with the resemblance which some of the forms depicted by 
Castellani bear to the breaking-down malarial parasites en- 
countered in spleen-puncture films from cases of chronic and 
relapsing malaria. 
Habitat. — Blood and spleen of a man : Ceylon. 




The H^MOSPOBiDiA are Coccidia-like forms specially modi- 
fied for parasitic life in the blood. There is alternation of 
hosts, asexual reproduction or schizogony taking place in the 
blood of Vertebrates and sexual reproduction or sporogony 
in the ahmentary canal of some blood-sucking Invertebrates. 
They are minute, usually intracellular parasites of red blood- 
corpuscles, showing motile amoeboid forms in their schizo- 
gonous cjT^cle in the Vertebrate host. Gametocytes and 
dimorphic gametes are formed, as in the Coccidia ; but the 
microgametes have no flagella as a rule and move like spiro- 
chetes, fertihzing a spherical macrogamete in the body of 
the Invertebrate host. The zygote is motile, and is known as 
an ookinete ; after becoming encysted it gives rise to a large 
number of naked sporozoites, which are introduced into the 
blood of a Vertebrate host. As they do not pass any stage 
of their hfe-history outside the body of a host, the sporozoites 
are not enclosed within a resistant membrane. 

Vertebrates of all classes — mammals, birds, reptiles, 
amphibians, and fish — are parasitized by different species. 
A number of them are known to occur in man and cause malaria, 
which works such havoc in India and other tropical countries. 
The Arthropods were their primary hosts, and they became 
secondarily introduced into the Vertebrates and adapted to 
a parasitic mode of hfe in the blood of the latter. 

The H^MOSPORiDiA are divided into four famihes. 

Identification Table of Families. 

1 (4). Parasites form hsemazoin pigment (with 

the exception of Leucocytozoon) 2. 

2 (3). Schizogony in the endothehal cells of the 

blood-vessels of internal organs of 

Vertebrates. Gametocytes in the peri- [Dofleiu, p. 210. 

pheral blood-corpuscles Hsemoproteidae 

3 (2). Schizogony in the peripheral blood of [p. 244. 

Vertebrates PlasmodiidaB Mesnil, 

4(1). Parasites do not form hsemazoin pigment. 5. 
5 (6). Schizogony in the endothelial cells of the 

blood-vessels of Vertebrates ; finally the 

parasites invade the red corpuscles, 

within which they occur as round, ovoid, 

rod-like or irregular forms. Show no [p. 294. 

tendency towards a paired arrangement . Theileriidse Du Toit, 


6 (5). Schizogony in the red blood-corpuscles of 
Vertebrates, with division into two or 
four ; of varying size and shape, and 

have a tendency to arrangement in [p. 301. 

couples of pear-shaped individuals Babesiidse Poche, 

1. Family H^MOPROTEID^ Doflein, 1916. 

Schizogony takes place in the endothehal cells of the blood- 
vessels of Vertebrates. Certain merozoites penetrate into the 
circulating red blood-ceUs, in which they develop into gameto- 
cytes. If the blood is taken up by a specific blood-sucking 
Invertebrate host, the gametoc3^es develop into gametes, 
which unite to form the zygotes, and the latter undergo 
sporogony, as in the family Plasmodiidse. 

Remarks. — It is important to remember that the schizogony 
cycle can only be observed in sections or smears from the 
internal organs of the host. In the absence of definite know- 
ledge regarding the occurrence of schizogony in the internal 
organs, the generic position can only be inferred from the 
morphology of the parasites themselves and their occurrence in 
relation to the nucleus of the infected corpuscle. In Hsemo- 
jproteus the organism is halter-shaped and grows round the 
nucleus without displacing it, whereas in Proteosoma (family 
Plasmodiidse) the nucleus of the red blood-corpuscle is ordinarily 
pushed to one side by the invading organism. 

Stiles (1925) gives " ookinete not known to encyst " as the 
character which serves to distinguish the family Haemoproteidse 
from the family Plasmodiidse. This statement appears to be 
based on the earlier work of Aragao. Helen Adie (1915) 
followed the complete development of the ookinete of 
Hssmoproteus columbas in the fly Lynchia maura. She 
confirmed that work in 1924, and Aragao (1927) also con- 
firmed it. Numerous oocysts were found in the wall of the 
stomach of the fly, and the various stages of development 
were found to resemble closely those of malarial parasites in 

Key to India Genera. 

1 (2). Young gametocytes enter the red blood- 

corpuscles ; when fully developed the 
gametocy te is halter-shaped, and produces 
pigment granules from the haemoglobin. . [Kruse, p. 211. 

2 (1) Gametocytes invade the immature red H^mopboteus 

blood-cells which have not yet produced 

haemoglobin, the host-cell is profoundly 

altered, becoming an elongated spindle, 

and the gametocyte does not produce [Danilewsky, p. 238. 

any pigment Leucocytozoon 


Genus H^MOPROTEUS Kruse, 1890. 
(8301. Haltbridium Labbe, 1894.) 

Hsemoproteus, Kruse, 1890, p. 371. 

Laverania (part), Grassi & Feletti, 1890, p. 463. 

Hsemamoeba (part), Grassi & Feletti, 1891, p. 463. 

Halteridium, Labbe, 1894, pp. 129, 151 ; 1899, pp. 78-9; Minchin 
1903, pp. 257-9, 267-9; Stephens & Christophers, 1904', 
pp. 319-20. 

Laverania (part), Laveran, 1899, pp. 603-6. 

Hxmocystidium, Castellani & Willey, 1905. pp. 84—5. 

Hxmoproteus, Minchin, 1912, pp. 365-9 ; Prowazek, 1912, pp. 566— 
88,; Castellani & Chalmers, 19J9, pp. 518-26; Wenyon, 1926 
pp. 885-902; Knowles, 1928, pp. 372-8; Reichenow,' 1929' 
pp. 972-9 ; Kudo, 1931, p. 288; Calkins, 1933, p. 566: Reiche- 
now, 1935, p. 375. 

The parasites grow in the endothelial cells of the blood- 
vessels of various organs into fairly large multi-nucleated 
schizonts, which then break up into very numerous merozoites. 
It is possible that some of them enter other endotheUal 
cells and again become schizonts, or, entering the red blood- 
corpuHcles, grow into gametocytes. They produce h»mozoin 
pigment granules at the expense of the haemoglobin of the 
host-cells. The fully-formed gametocyte encircles the nucleus 
of the red blood-corpuscle Hke a halter (hence the name 
Halteridium Labbe) and usually does not force it out of place. 
Parasitic in birds and reptiles. Sporogony takes place in the 
body of an Arthropod. 

Remarks. — Danilewsky (1889) was the first to record the 
parasites referable to this genus. Labbe (1899) considered 
that the halteridia of different birds belong to one species, 
to which he restricted the name H. danilewskyi (Grassi & 
Feletti). They are now known from hundreds of dififerent 
species of birds, and many species are recognized. They have 
often been confused in the past with another pigmented parasite 
of the blood of birds which belongs to the same family as the 
human malarial parasite. Researches of Danilewsky, Laveran, 
Kruse, Grassi, Feletti, and others have shown the difference 
between Haemoproteus {Halteridium) and Proteosoma, viz., that 
schizogony takes place in the endothehal cells in the former 
and only gametocytes are found in the red blood-corpuscles, 
while in the latter schizogony takes place in the red blood- 
corpuscles and thus both merozoites and gametocytes can be 
found in them. 

It was in the Hsemoproteus in the blood of the crow that 
MacCullum (1897) demonstrated for the first time that the 
so-called " flagellating body " was the male gametocyte, 
and the process of ex-flagellation produced the male gametes, 
which eventually fertilized female gametes and led to the 
formation of motile zygotes or ookinetes. 



Coatney (1936) has catalogued 45 species of Hmmoproteus 
and has given a complete hst of 650 avian and 22 reptiUan 
hosts from which they had been recorded up to the time 
of his compilation. 

147. HaBmoproteus antigonis de Mello. 

■fHsemoproteus antigonis, de Mello, 1935 6, p. 471 ; 1937 a, p. 100. 

Female gametocyte with alveolar cytoplasm, stained blue 
with Lieshman's stain ; nucleus compact, oval, shghtly 
subcentral, stained rose, lodged in a vacuole. Male gametocyte 
pale violet ; nucleus granular, central. Pigment coffee-brown, 
scattered over the body ; in the female gametocyte often in 
clusters or in two large masses united by a batonette. 

Remarks. — The parasite may possibly be the same as that 
recorded by Scott (1926) from Antigone antigone (Linn.). 

Habitat. — Blood of Anthropoides virgo (Linn.) : Pobtfguesb 
India, Junagad. 

148. Hsemoproteus asturis de Mello. 

■fHsemoproteus asturis dussumieri, de Mello, 1935 b, p. 469 ; 1937 a, 
p. 100. 

Female gametocytes with alveolar cytoplasm, stained light 
TdIuc with Leishman's stain, nucleus pale rose. Male gameto- 
cyte colourless, nucleus more conspicuous. General form 
oval, especially in male, rarely halter-like in female. Nucleus 
sometimes subcentral. Pigment granules dark brown, of 
•different sizes, irregularly scattered in both kinds of gameto- 

Remarks. — De Mello has given to this form a trinomial 
designation to avoid confusion with the unnamed Hsemoproteus 
of Astur plumbarius recorded by Wasielewsky in Germany 
in 1908. 

Habitat. — Blood of Astur badius dussumieri (Temm. & 
Lang.) : Portxjgtjesb India, Corlim (Ilhas). 

149. Hsemoproteus cerchneisi, sp. nov. 

■\H3emoproteus sp., de Mello, 1935 b, p. 472 ; 1937 a, p. 100. 

Female gametocyte halter-like, cytoplasm alveolar, stained 
greyish-blue with Leishman's stain ; nucleus generally round 
and compact, but often quadrangular and consisting of a central 
mass with appendages all round, subcentral on the convex 
border of the parasite, stained hght red and lodged in a vacuole : 
pigment coffee-brown with oHve tone, granules or clusters 
irregularly distributed. Male gametocyte oval or round, 
and causing a sort of hernia on the periphery of the red cell ; 
cytoplasm colourless or light violet ; nucleus subcentral or 


central, small, elliptic or irregular, containing one chromatic 
dot fairly visible, surrounded by the rest of the chromatic 
substance which is hardly visible, stained pale rose ; pigment 
coffee-brown with oHve tone, situated at the poles. 

Remarks. — Unnamed species of Hsemoproteus have previously 
been recorded from several species of Cerchneis in different 
parts of the world. According to de Mello, Wasielewski and 
Wiilker (1918) recorded the parasite of this bird in Europe as 
H. danilewskyi var. tinnunculus, and not having access to 
that paper he has refrained from giving his form a specific 
name. On comparing his description with Wasielewski and 
Wiilker 's figure, as reproduced by Reichenow, the nucleus 
of the microgametocyte in var. tinnunculus is large and does 
not correspond with the description as given by de Mello. 
I have, therefore, named the parasite found by de Mello as 
H. cerchneisi. 

Habitat. — Blood of Cerchneis tinnunculosus objurgatus 
Stuart Baker : Portuguese India, Pragana. 

150. Hsemoproteus columbae CelH & San FeHce. (Figs. 104, 

Hsemoproteus danilewskii (part), Kruse, 1890, p. 359. 

Hsemoproteus columbse, Celli & San Felice, 1891, pp. 517-18, 541-8, 
pi. vi; Ed. & Et. Sergent, 1907, pp. 264-74, pis. vi, vii. _ 

Halteridium columbse, Aragao, 1908, pp. 154r-67, pis. xi-xiii. 

Haemcproteus columbse, Minchin, 1912, pp. 365-6, fig. 157. 

Hsemoproteus sp., Sergent & Beguet, 1914, pp. 21-3. 
■\Heemoproteus sp., Acton & Knowles, 1914, pp. 663-90, pis. xlvii-li ; 
Alcock, 1914, p. 584 ; Helen Adie, 1915, pp. 671-80. 

Hsemoproteus columbse, Castellani & Chalmers, 1919, pp. 525-6, 
figs. 185, 186; Helen Adie, 1924, pp. 605-13, 2 pis.; 1925, 
pp. 9-15, 5 figs. ; Wenyon, 1926, pp. 886, 888-96, 1314, 
figs. 383-6 ; Knowles, 1928, pp. 372-7, figs. 87, 88 ; Reichenow, 
1929, pp. 973-5, 978, figs. 938-40, 942 ; Kudo, 1931, p. 288, 
fig. 122e, /; Ruiz Martinez, 1934, pp. 96-8; Coatney, 1936, 
p. 88 ; de Mello, 1937 a, p. 99. 

Schizogony takes place in the endothehal cells of the blood- 
vessels of various organs of the pigeon, particularly the lungs. 
The youngest schizonts are minute cytoplasmic bodies with 
a single nucleus within the cytoplasm of an endothehal ceU 
(fig. 104, P) . Growth, nuclear multiphcation, and segmentation 
into fifteen or more small uninucleated unpigmented masses 
take place (fig. 104, Q). Each of these cytomeres grows; its 
nucleus undergoes repeated division till the host-cell, now 
considerably hypertrophied, is filled by a number of multi- 
nucleate bodies, each of which is surrounded by a fine membrane. 
Within this membrane the multinucleate cytomere divides 
into an enormous number of merozoites (fig. 104, R-W). The 
endothelial cell finally breaks down and the merozoites escape 
into the blood-stream. It is possible that some of them 

Fig. 104. — Life-cycle of Haemoproteus columhse Celli & San Felice. The 
stages above the dotted line occur in the pigeon, those 
below in the fly. A^-C-^, growth of female gametocyte 
in red blood-corpuscle ; A^—G^, growth of male gameto- 
cyte ; D^, E-^, rounding off of female gametocyte and 
escape from cell ; D,^, E^, rounding off of male gameto- 
cyte and formation of male gametes ; F, fertiUzation ; 
6—L, formation of ookinete, which finally makes its 
way through the stomach-wall of Lynchia maura ; 
M, young oocyst on stomach-wall of Lynchia maura ; 
N, mature oocyst filled with sporozoites, which eventually 
enter the salivary glands of the fly and are thence injected 
into the pigeon ; O, sporozoite entering an endothelial cell 
of a blood-vessel of the pigeon ; P, growth of the sporozoite 
in a mononuclear cell ; Q, primary schizogony into a niunber 
of uninucleate bodies ; R—V, each uninucleate body increases 
in size and becomes multinucleate ; W, segmentation into 
numerous minute young gametocytes which enter red 
blood-corpuscles. (From Wenyon, after Aragao and Adie.) 



■enter other endothelial cells and again become schizonts. 
Others enter the red blood- corpuscles and are seen as minute 
cytoplasmic bodies with a single nucleus and often a vacuole 
(fig. 104, A-j^, A^). Sometimes as many as a dozen young forms 
may be present in a single corpuscle, and thus in dry films, 
by an approximation and overlapping of the parasites, falsely 
suggest a schizogony. The corpuscles with such multiple 
infections generally die, so that it is rare to find even two 
fully developed gametocytes in a single corpuscle. The young 

I'ig. 105. — Development of Heemoproteus columbse Celli & San Felice on 
stomach- wall of Lynchia maura. A, flattened stomach 
showing numerous oocysts of various sizes ( X c. 72) ; 
B, edge of stomach more highly magnified, showing 
mature and immature oocysts with pigment granules 
( X c. 450) ; G, intact oocyst and ruptured oocyst, with 
discharged sporozoites and residual cytoplasm { X c. 600) ; 
D, free sporozoites. (From Wenyon, after Adie.) 

gametocyte becomes elongate, and granules of brown or black 
pigment appear in the cytoplasm (fig. 104, B, C) : it grows 
roimd one side of the nucleus, pushing it to one side of the 
oell, but the latter retains its original shape and size. The 


fully developed gametocyte is of an elongate sausage-shape^ 
and almost completely encircles the nucleus : it shows a central 
nucleus and a number of pigment granules distributed through 
its cytoplasm. Gametocytes are distinguishable as male and 
female. Male gametocyte possesses a hyahne cytoplasm, 
staining a pale blue or pinkish colour with Romanowsky 
stains, and a rather larger nucleus consisting of a membrane 
enclosing a number of fine chromatin granules (fig. 104, B^, C^). 
Female gametocyte has a denser cytoplasm, stains more deeply 
blue, and possesses a more compact nucleus (fig. 104,5^, C^). 

Further development of the gametocytes takes place in the 
fly, Lynchia maura. The male gametocyte produces male 
gametes by exflagellation, the female gametocyte gives rise 
to a single female gamete, and fertilization produces a zygote, 
which is a motile vermicule or ookinete. The ookinetes 
penetrate the hinder portion of the mid-gut of the fly and 
produce pigmented oocysts on the outer surface of the wall. 
The mature oocyst (fig. 105, B) measures about 36 jx in diameter, 
and gives rise to a very large number of sporozoites (fig. 105, 
G, D), measuring up to 10 /x in length. These are set free by 
the rupture of the cysts, invade the salivary glands of the 
fly, and are reintroduced into the body of the pigeon. 

Remarks. — Acton and Knowles (1914) studied the develop- 
ment in the pigeon and concluded that schizogony takes place 
only in the lung, but other observers have shown that it takes 
place in the endothehal cells of the blood-vessels of other organs 
as well. Clean pigeons exposed to infection by infected flies 
show young gametocytes in the blood in about four weeks, 
during which schizogony takes place a number of times. 

Ed. and Et. Sergent were the first (1906) to transmit the 
infection experimentally to pigeons in Paris by means of 
infected flies {Lynchia maura), received from Algiers. The 
asexual cycle as occurring in the endothelial cells was first 
described by Aragao (1908). Helen Adie (1915), working in the 
Punjab, was the first to follow the sporogony of the ookinetes in 
L. maura, and later, working at Algiers (1924, 1925), confirmed 
her previous work in India. According to her researches, 
flies which have hved on infected pigeons for ten to twelve days, 
during which they fed daily, showed all stages of development 
of the parasite from the ookinete to the sporozoites in the 
salivary glands. 

De Mello and de Sa (1916) have described a process of 
schizogony in Hsemoproteus columhse taking place in schizonts 
which were originally in the red blood-corpuscles and then 
became free in the plasma. If their observations were correct, 
they must have been dealing with a species of Proteosoma. 
These observations are recorded under Proteosoma cohimbse 
(de Mello & de Sa). 



Habitat. — Blood of the oommon pigeon, Golumha livia 
Gmelin, and body of Lynchia maura Bigot : Punjab, Amballa ; 
Bengal, Calcutta. 

151. Haemoproteus coraciae de MeUo & Afonso. (Fig. 106.) 

"fHcetnoproteus coraciae benghalensis, de Mello & Afonso, 1935,. 
pp. 67-8, pi. i ; de Mello, 1937 a, p. 100. 

Sexual dimorphism of gametocytes shown by tinctorial 
reactions of cytoplasm, which stains blue in female gameto- 
cytes and is colourless or shghtly yellowish-blue in male 
gametocytes. Young female gametocytes small, more or less 
ovoid ; when full grown are typical halteridia, embracing the 





Fig. 106. — -Hsemoproteus coracise de Mello & Afonso. A-C, female 
gametocytes ; D, free female gametocyte ; E-G, male 
gametocytes ; H, free male gametocyte. (After de Mello 
and Afonso.) 

nucelus of the red cell and displacing it to the periphery. 
These halteridia may be pointed, with a tail-like appendage, 
but when full grown they are broad, regular, lodging the 
nucleus of the host-cell in their concavity ; cytoplasm alveolar, 
not staining uniformly, the blue colour more pronounced at 
the poles and hghter in the centre, and showing violet rings in 
some specimens ; nucleus compact or more or less granular ; 
pigment yellow- brown, in minute granules or big dots, showing 
a tendency to collect at the poles. When free the female 
gametocytes are roundish or oval, with blue cj^oplasm, 
nucleus compact or irregular, pigment irregularly scattered over 



the body in clusters or isolated granules. Male gametocytes 
show the same form as the female, a Httle more irregular 
in young stages ; nucleus in the form of an irregular spireme. 
Pigment has the same appearance, but is more definitely 
collected at the poles. When free the male gametocytes 
are round, the cytoplasm showing a slight violet- bluish tone. 
Infected cells are slightly hypertrophied. 

RemarTcs. — Hwrnoproteus has been previously recorded from 
Coracia indica Linn, by Plimmer (1912, 1914) and from other 
species of Coracia from other parts of the world, but none of 
these appears to have been given a specific name. De Mello 
rejects the idea that all Hsemoproteids from birds in general 
are identical with H. daniUwskyi. 

Habitat. — Blood of Coracia benghalensis benghalensis Linn. : 
PoRTtTGUBSE India, CorHm (Ilhas). 

152. Haemoproteus corvi, nom. nov. (Fig. 107.) 

•\Haemoproteus sp., Donovan, 1904; Castellani & Willey, 1905, 
p. 385, pi. xxiv, fig. 5 ; p. 400 ; Dobell, 1910, p. 71. 

'fHsenioproteus du Corvus macrorhynchus, de Mello, de Sa, de 
Sousa, Dias, & Noronha, 1917, pp. 10-11, pi. i, figs. 45-71 ; 
de Mello, 1937 a, p. 100. 


Fig. 107. — Hsemoproteus corvi, nom. nov. A, female gametocyte ; 
B, male gametocyte ; G, so-called "Acton body," showing 
schizogony. (After de Mello, de Sa, de Sousa, Dias, and 

Young parasites small, lanceolate. Female gametocji^e 
halter-shaped, stained blue by Leishman's stain. Nucleus 
a large chromatic mass, stained deep red, situated usually 
in the concavity of the halteridium, nearer one pole than the 
other, and quite close to the nucleus of the infected corpuscle ; 
pigment collected at one pole, but in the fully developed 
gametocyte may occur at both poles. Male gametocyte ovoid 
or halter-shaped, stained but Httle, almost yellowish ; nucleus 
contains two or three chromatin granules, and is situated 
near one pole, while the pigment is situated near the other. 


When fully developed the nucleus may be vesicular or compact 
and linear ; sometimes the pigment may occur near both the 
poles also. Schizogony takes place in the lung and not in 
the peripheral blood. It is very similar to schizogony as 
described in H. columbse. In the intracorpuscular phase the 
schizonts are small, medium-sized or large, and in the extra- 
corpuscular phase the " Acton body " shows a single nucleus 
or, as the result of repeated division, forms a number of nuclei 
leading to the formation of as many merozoites. 

Remarks. — De Mello and his colleagues referred the parasite 
to the genus Hsemoproteus, but described schizogony as taking 
place in "Acton bodies " that have come out of the blood- 
corpuscle. As, however, they state that schizogony takes place 
in the lung (presumably in the blood-vessels), it is conceivable 
that the so-called "Acton bodies " may have been endothelial 
cells of the blood-vessels that are being carried along in the 
plasma. The form is consequently retained in the genus 
Hasmoproteus pending further investigation. 

Habitat. — Blood and smears from the lung of the crow, 
Corvus levaillanti macrorjiynchus Blanf. & Gates : Ceylon ; 
PoETFGTJESE India, Nova Goa ; and the Indian house-crow, 
Corvus splendensYi&m. : India (locahty not cited by Donovan) ; 
Ceylon ; Portuguese India, Nova Goa. 

153. Hsemoproteus danilewskyi (Grassi & Feletti). (Fig. 108.) 

Haemoproteus (part), Kruse, 1890, p. 359. 

Laverania danilewskyi, Grassi & Feletti, 1890, p. 463. 

Halteridium danilew&kyi, Labbe, 1899, p. 79, fig. 145 ; Minchin, 

1903, pp. 266, 268, 269, 347, 348, 349. 
■fHalteridium danilewskyi, Stephens & Christophers, 1904, pp. 319- 

21, fig. 67 ; Castellani & Willey, 1904, pp. 83-4, figs. 7-9. 
Hsemoproteus danilewskyi, Minchin, 1912, p. 365. 
'fHcemoproteus danilewskyi, Plimmer, 1912, pp. 416, 417; 1913, 

p. 148; 1914, p. 189; 1915, p. 130 ; 1916, p. 85 ; 1917, p. 32. 
Hsemoproteus danilewskyi var. tinnunculus, Wasielewski & Wiilker, 

1918, p. 115. 
Hsemoproteus sp., Wenyon, 1926, p. 1383. 
Hsem,oproteus danilewskyi, Reichenow, 1929, pp. 975-7, figs. 943-6; 

Coatney, 1936, p. 88. 
'\ Hsemoproteus sp., de Mello, 1937 a, p. 99. 

Young oval stages of the trophozoite present, though not 
common. Two kinds of fully developed gametocytes (referred 
to as trophozoites by the discoverers) present in approxi- 
mately equal numbers. The female gametocytes are stained 
distinctly blue with Leishman's stain, leaving a clear tract in 
the centre, and with pigment granules scattered more or less 
throughout the cytoplasm. The male gametocytes are shorter 
and stouter, appearing nearly white or very faintly bluish- 
white, owing to greater density of cytoplasm, and the pigment 



granules are aggregated at the two ends. The male becomes 
shorter, thicker, and finally nearly round. In one instance 
a double infection of a blood- corpuscle by the two kinds of 
gametocytes was seen. 

Remarks. — Wasielewski and Wiilker (1918) described from 
the kestrel a parasite which they named H. danilewskyi var. 
tinnunculus, accepting the view that many species of birds can 
harbour the same species of parasite. They point out that in 
kestrels the infection is either acute or chronic. Acute 
infection occurs in young birds, which are heavily infected 
and show schizonts in the various organs. When this subsides 
chronic infection supervenes and is characterized by relapses 

Fig. 108. — Hsemoproteus danilewskyi var. tinnunculus Was. & Wiilk. 
A, female gametocyte ; B, male gametocyte ; C, double 
infection with both types of gametocytes. (After Wasielew- 
ski and Wiilker.) 

of a milder type extending over several years. Different 
stages in the life-cycle are fully described by them. 

Habitat. — Blood of the scops owl, Otus hakkamcena hakka- 
moena Pennant : Ceylon, Colombo. Also in the blood of the 
following birds from India in the Zoological Gardens, London : — 

Anas {Fuligula) baeri Radde (Bayer's pochard). 
Kittacincla macroura Blanf. & Gates (shama). 
Copsychus saularis (Linn.) (Indian dial-bird). 
Cor acta benghalensis indica Linn. (Indian roller). 
Garrulax albigularis Gould (white-throated jay-thrush). 
Glareola pratincola Linn, (pratincole). 
Garrulus lanceolatus Vigors (jay). 

Melophus melanicterus (Gmelin) (crested black bunting). 
Mesia argentauris Hodgs. (silver-eared babbler). 
Nettapus coromandelianus GmeHn (cotton-teal). 
Propasser rhodochrous (Vigors) (finch). 

Prunella strophiata jerdoni Brooks {=Tharrhaleus jerdoni 
Blanf. & Gates) (Jerdon's hedge-sparrow). 


154. Hsemoproteus glaucidii de Mello. 

■\Hsemoproteus glaucidii, de Mello, 1935 b, p. 473 ; 1937 a, p. 100. 

Female gametocyte vacuolated, stained deep blue with 
Leishman's stain ; nucleus spherical and central or elongate 
and situated on the convex border, stained rose ; pigment 
scattered. Male gametocyte almost unstained ; nucleus large, 
without definite borders, containing irregular chromatic 
masses, subcentral in halter-like forms, central in the rounded 
ones. Infected red cell hypertrophied, with its nucleus 
retaining its position when the corpuscle contains a female 
gametocyte, and displaced when there is a male gametocyte. 

Remarks. — De Mello thinks that perhaps the parasite is the 
same as the unnamed Hsemoproteus of Glaucidium perlatum 
recorded by A. & M. Leger (1914) in Niger, or perhaps it is 
only a new variety of H. noctuse CelH & San Felice (1891). 

Habitat. — Blood of Glaucidium radiatum Tickell : Portu- 
OTJESE India, Canacona. 

155. Haemoproteus gymnorhidis de Mello. 

fHeemoproteus sp., Plimmer, 1913, p. 148. 

•fHmmoproteiis gymnorhidis, de Mello, 1935 6, p. 474 ; 1937 a, p. 100. 

Female gametocytes halter- shaped, very slender, larger 
at the poles, cytoplasm alveolar, stained deep blue with 
Leishman's stain, more deeply at the poles ; nucleus central 
or subcentral, rod-like, oval or bilobed ; pigment in clusters 
of large granules at the centre and at the poles, in the latter 
situation sometimes as a fine dust, rendering more intense the 
dark staining of the cytoplasm in this part. The pigment 
spreads a little round the granules, diffusing into the cytoplasm. 
Male gametocyte halter-shaped or oval, not stained or shghtly 
blue ; nucleus as in the female gametocyte. Pigment granules 
very minute, mostly at the poles. Infected red cell hyper- 

Remarks. — The parasite is probably the same as the unnamed 
Haemoproteus from the Indian Gymnoris flavicollis recorded by 
Plimmer (1913), as Gymnoris flavicollis is a synonym of 
G. xanthocollis. 

Habitat. — Blood of the yellow-throated finch, Gymnoris 
xanthocollis Burton : Portuguese India, Pragana ; also 
from a specimen of the same species in the Zoological Gardens, 

156. Hsemoproteus herodiadis de Mello. (Fig. 109.) 

■\Hsemoproteus herodiadis, de Mello, 1935 a, pp. 351-2, pi xlii 
fig. 2 ; 1937 a, p. 101, pi. 1, fig. 1. 

Young stages roundish oval, fully developed halteridia never 
embracing the nucleus of the red cell. Fusiform or irregularly 



bent forms also occur. Sexual dimorphism of the gametocytes 
only visible in the tinctorial reaction of the cytoplasm to 
Romanowsky stains, it being light blue and alveolar in female 
gametocytes and colourless, almost white, in male gametocytes. 

Fig. 109. — Heemoproteus herodiadis de Mello. 
A, female gametocyte ; B, male gametocyte. (After de Mello.) 

Brownish-black pigment irregularly distributed in both. 
Infection heavy. 

Remarhs. — De Mello thinks that the form is perhaps similar 
to that found in Herodias alba in the Belgian Congo by 
Rodhain, Pons, Vandenbranden, and Bequaert (1913). 

Habitat. — Blood of the heron, Egretta intermedia intermedia 
(Wagler) : Porttjgfbsb India, lake of Carambolim. 

157. Haemoproteus kopki (de Mello). (Fig. 110.) 

'\Hsemocystidium kopki, de Mello, 1916, pp. 8-10, pi. i, figs. 1-17 ; 

de Mello, de Sa, de Sousa, Dias, & Noronha, 1917, p. 13. 
Hsemoproteus simondi, Wenyon, 1926, p. 902. 
^Hsemoproteus kopki, de Mello, 1934 a, p. 378; 1934 6, pp. 1785- 

99, pi. i. 
Hsemoproteus kopki, Coatney, 1936, p. 88. 

Yoimg stages round and non-pigmented. Female gameto- 
cyte variable in form, usually halter-shaped or oval ; cyto- 
plasm finely granular, staining deep blue with Leishman's 

A B C D 

Fig. 110. — Hsemoproteus kopki (de Mello). 

A, B, female gametocytes ; C, D, male gametocytes. 
(After de Mello.) 



or May-Grunwald Giemsa stain, with small irregular alveoli 
uniformly distributed all over the body, sometimes with larger 
vacuoles or even larger clear zones ; nucleus a compact 
chromatic granule, stained faintly rose ; pigment granules very 
small, and scattered irregularly all over the body. Male 
gametocyte usually ovoid, not halter-shaped ; cytoplasm, 
not staining or only staining light straw-yellow or light grey, 
with very distinct vacuoles, rarely without them ; nucleus 
granular, compact, sausage -shaped or oval, only faintly 
staining, and always larger than in the female gametocyte ; 
pigment granules larger than in the female gametocjrfce, and 
situated in one, two or three well circumscribed vacuoles. 
Infected red ceU hypertrophied, dehsemoglobinized, with its 
nucleus displaced. Schizogony in the monocytes or endotheUal 
cells of the lung. 

Eemarks. — ^De Mello (1934 b) has described the schizogony 
occurring in the pulmonary epithelium of the infected Uzard 
as very similar to that of Hsemoproteus in birds. This, 
along with similarity in other characters, definitely estabhshes 
the identity of Hsemocystidium, with Hsemoproteus. He has 
further shown that H. kopM is specifically distinct from 
H. simondi, and not identical as was believed by Wenyon. 

Habitat. — Blood and lungs of Hemidactylus brooki Gray : 
Portuguese India, Nova Goa. 

158. Hsemoproteus machlolophi de Mello. (Fig. 111.) 

^Hsemoproteus danilewskyi, Plimmer, 1912, p. 417. 
^Hsemoproteus machlolophi, de Mello, 1935 a, pp. 353-4, pi. xliii, 
fig. 3 ; 1937 a, p. 100. 

Halteridia forms not very conspicuous ; nucleus sub- 
central, stained pale rose with Leish man's stain. Female 

A B 

Fig. 111. — Hsemoproteus machlolophi (de Mello). 
A, female gametocyte ; B, male gametocyte. (After de Mello.) 

gametocytes very irregular, denticulated in outline, stained 
greyish-blue ; pigment dark-brown, granules irregularly 
scattered in the cytoplasm. Male gametocytes of a regular 



outline, stained clear, almost white ; pigment dark-brown, 
granules, showing a tendency to collect at the poles. 

Habitat. — Blood of the yellow- cheeked tit, Machlolophus 
xanthogenys (Vigors) : from India, in the Zoological Gardens, 
London ; also from Portuguese India, Siroda (Ponda). 

159. Hsemoproteus metchnikovi (Simond). 

(Fig. 112.) 

150-2; 19016, 

■^Hsemamoeba metchnikovi, Simond, 1901 a, pp. 

pp. 338-43, pi. viii, figs. 1-20. 
Hsemocystidium metchnikovi, Castellani & Willey, 1905, pp. 84—5 ; 
Castellani & Chalmers, 1919, p. 516 ; Wenyon, 1926, pp. 899, 
901, 1396 ; Knowles, 1928, p. 378 ; Reichenow, 1929, p. 978 ; 
Coatney, 1936, p. 89. 

Gametocytes rarely exceeding half the blood- corpuscle in 
size, with a small number of pigment granules, and not causing 

Eig. 112. — Hsemoproteus metchnikovi (Simond). A, tailed non-pig- 
mented form ; B, large non-pigmented vermicule form ; 
C, young pigmented form ; D, female gametocyte ; E, female 
gametocyte and male gametocyte in the same corpuscle ; 

F, fully developed female gametocyte showing nucleus ; 

G, leucocyte from the spleen showing two female and 
one male gametocytes; H, leucocyte from the lung showing 
male gametocyte. (After Simond.) 

a displacement of the nucleus or distortion of the corpuscle. 
In some cases a blunt prolongation of the parasite extends 
up the side of the nucleus, but typical halteridia forms are 
not seen. Male gametocytes faintly staining, containing 
larger nucleus and coarse irregularly distributed pigment ; 
female gametocytes deeply staining, containing smaller nucleus 
and finer pigment. Sometimes two gametocytes are found 
in the same corpuscle. 

Dimensions. — Gametocytes 6-10 /a in diameter. 



Remarks. — Simond (1901 a, b, e) described two distinct 
species from the same host, one non-pigmented and hsemo- 
gregarine-like, the other pigmented hsemamoeba-hke. Heemo- 
gregarina hankini is very rare, but shows the tailed and vermi- 
cule forms similar to the Hsemogregarines of other tortoises. 
Hsemoproteus metchinikovi shows sexual differentiation, the 
faintly staining forms with coarse grains of pigment being 
regarded as male and the deeply staining with finer grains 
of pigment as female gametocytes. 

Habitat. — Blood of the tortoise, Ghitra indica (Gray) 
{=Trionyx indicus) of the Ganges and the Jumna : United 
Provinces, Agra. 

160. Hsemoproteus orioli de Mello. 

■fHsemoproteus orioli, de Mello, 1935 b, p. 469 ; 1937 a, p. 100. 

Female gametocyte stained Hght blue with Leishman's 
stain ; nucleus central or sub central, constituting a more 
or less compact chromatic dot. Male gametocji^e whitish ; 
nucleus central or subcentral, composed of an irregular spireme. 
Pigment Hght brown, granules of different sizes irregularly 
distributed, sometimes situated near the poles in both sexes. 

Remarks. — De Mello thinks that possibly the parasite may 
be identical with the unnamed H£em,oproteus of Oriolus 
Sagittarius recorded by Cleland and Johnston in AustraHa in 
1912, and of 0. galbula recorded by Cardamatis in Greece 
in 1919. 

Habitat. — Blood of Oriolus oriolus kundoo Sykes : Portu- 
guese India, Nova Goa. 

161 . Hsemoproteus raymundi de MeUo & Raimundo. (Fig. 1 13.) 

■\Heeinoproteus raymundi, de Mello & Raimundo, 1934 a, pp. 97-9, 
pis. xii, xiii, text-fig. 1 ; 1934 b, pp. 1437-40, pi. i ; 1937 a, 
pp. 98, 100. 

Free trophozoites, uni- or multinucleated, are found in the 
lung, giving rise to schizonts, rosettes, and merozoites. The 

Fig. 113. 



-Hsemoproteus raymundi de Mello & Raimundo. A, female 
gametocyte ; B, male gametocyte ; C, schizogony in the 
lung. (After de Mello and Raimundo.) 


schizogony cycle is particularly simple. The merozoites remain 
free among the blood-corpuscles or the cells of the host-tissue. 
Sections of lungs and spleen also show schizonts withia the 
red corpuscles and free merozoites among them. In the blood 
the red corpuscles show halteridia or reniform gametocytes. 
Female gametocyte stained deep blue with Leishman's stain ; 
cytoplasm finely vacuolated ; nucleus eUiptical ; pigment 
granules deep brown and slightly larger than in the male, 
Male gametocyte stained Hght blue ; nucleus granular ; 
pigment granules finer. Gametocytes may become extra- 
cellular and free in the plasma, and assume an oval or rounded 

Dimensions. — Gametocytes 6-8 jtt by 2'5-3/x ; infected cor- 
puscles 8-9 ju, by 4-4-5 /x. 

Remarks. — ^When the gametocytes are full grown the para- 
sitized corpuscles become hypertrophied, decolorized, and 
sometimes reduced to a thin border roimd the parasite. The 
nucleus of the corpuscle is dislocated to the periphery, and 
finally disappears. 

Habitat. — Peripheral blood and sections of the lung and the 
spleen of Leptocoma zeylonica (Linn.) : Portuguese India, 
Nova Goa. 

162. HsBmoproteus rileyi Malkani. (Fig. 114). 

'^Hsemoproteus rileyi, Malkani, 1936, pp. 155-7, pi. iv. 

Gametocytes show a variety of form, such as rings, pear- 
shaped or elongate forms, spindles, boomerangs, and typical 

Fig. 114. — Haemoproteiis rileyi MsAak.axii. (After Malakani.) 

halteridia encirchng the nucleus of the infected red blood- 
corpuscles. Male gametocytes stained a pale blue with 
Leishman's stain ; nucleus central, large, consisting of a number 
of chromatin granules stained intensely red enclosed by a mem- 
brane ; brownish-black pigment granules collected together. 
Female gametocytes stained more deeply and the nucleus 
not granular but compact. The number of gametocytes in 
each corpuscle varied from one to four. In some there was 
a mature gametocyte on each side of the nucleus, both of the 
same sex or opposite sexes, No schizonts could be seen in 
the blood. 


Remarks. — The parasite is believed to be responsible for 
a serious illness which ended fatally. 

Habitat. — Red blood- corpuscles of the peacock, Pavo 
cristatus Linn. : Bihak, Patna. 

163. Haemoproteus simondi (Castellani&Willey). (Fig. 115). 

■\Hsemocystidium simondi, Castellani & Willey, 1904, pp. 84-5, 
figs. 10-16; Robertson, 1908, pp. 181-2; Dobell, 1910 a, 
pp. 68, 69 ; 1910 b, pp. 123-32, pi. vii ; Castellani & Chalmers, 
1919, p. 516, fig. 173. 
Hsemoproteus simondi, Wenyon, 1926, pp. 899, 902, 1388 ; Knowles, 
1928, p. 378 ; Reichenow, 1929, pp. 978-9, fig. 949. 
■\Hsemoproteus simondi, de Mello, 1934 a, pp. 6-10, 14-15, pi. ii. 
Heeinoproteus simondi, Coatney, 1936, p. 89. 

Trophozoite small, rather irregular or amoeboid, with a zone 
of pigment granules across the centre, at first only sHghtly 
displacing the nucleus of the corpuscle ; by its further growth 
the nucleus of the corpuscle is more displaced. Sometimes 


Pig. 115. — Hsemotr opens simondi (Castellani & Willey). A, female 
gametocyte ; JB, male gametoeyte ; C, double infection 
with a male and a female gametocyte. (After Dobell & 

the parasite is round, lenticular or oval, the oval form nearly 
filhng the corpuscle and moulding itself on the nucleus of the 
blood-cell. Spherical or discoidal forms are gametocytes. 
In the male type, body faintly granular ; stains a delicate 
pale blue with Leishman's stain, and possesses numerous small 
pigment granules scattered round the periphery. In the female 
type, body stains dark blue ; pigment granules, though 
numerous, are slightly larger, and a varying number of 
vacuoles always present. 

Dimensions. — Gametocytes about 18/x, in length by 9 /i in 

Remarks. — Dobell (1910) describes schizonts in the red 
blood-corpuscles. These are round bodies about 8^ in 
diameter, and are supposed to divide into two or four mero- 
zoites : but probably this apparent division was an artefact 
due to fusion of adjacent parasites during the drying of the 



blood-films, as shown by Wasielewski and Wtilker in the case of 
the H^moproteus of the kestrel. De Mello (1934) has carefully- 
compared his observations on H. kopki with the original de- 
scriptions of H. simondi, and has come to the conclusion that 
the two are distinct, and not identical as was supposed 
by Wenyon. The chief characters in which the two differ 
are as follows : — InH. simondi the male gametocyte is without 
vacuoles and with pigment granules finer than in the female 
and scattered over the body, whilst in H. kopki it is without 
distinct vacuoles ; the pigment granules are generally larger 
than in the female and are collected in vacuoles. As regards 
the female gametocyte, this has numerous vacuoles and the 
pigment granules are larger than in the male in the former 
species ; but is without definite vacuoles and the pigment 
granules are finer than in the male in the latter species. 

Habitat. — Blood of Hemidactylus leschenaulti Dum. & Bibr.: 
Ceylon, Trincomalee ; Portfgijese India, Mamadu, near 

164. Hsemoproteus sturni de Mello. 

■fHxmoproteus sturni, de Mello, 1935 6, pp. 473-4 ; 1937 a, p. 100. 

Female gametocyte vacuolated, stained Hght blue with 
Leishman's stain ; nucleus ovoid, central or subcentral ; 
pigment absent or irregularly scattered. Male gametocyte 
halter-like or oval, almost unstained ; nucleus conspicuous, 
compact, central or subcentral ; pigment at the poles. 

Remarks. — ^De Mello thinks that the parasite may perhaps 
be the same as the unnamed H^moproteus of Sturnus vulgaris 
Linn, recorded by CeUi and San Felice (1891) in Italy, by Labbe 
(1894) in France, by Wasielewski (1896) in Germany, and by 
Coles (1914) in England. 

Habitat. — Blood of Sturnia malabarica (GmeHn) : Portu- 
guese India, Ponda. 

165, Hsemoproteus upupse de MeUo. 

■fHsemoproteus upupse, de Mello, 1935 b, p. 472 ; 1937 a, p. 100. 

Female gametocyte stained bluish, but not uniformly, with 
Leishman's stain ; nucleus compact, spherical, ovoid or 
ribbon-like, subcentral, stained deep rose ; pigment in granules 
or clusters, irregularly distributed. Male gametocyte colour- 
less, with the borders having a very slight bluish tinge ; nucleus 
large, granular, pseudospirematic, the threads being irregularly 
interwoven or disposed in parallel lines obhquely to the long 
axis of the parasite, stained pale rose or violet-rose ; pigment 
granules small, often dust-like, and never fused together, 
scattered over the body or at the poles, fewer than in the 
female gametocyte. 


Remarks. — The parasite may possibly be the same as that 
recorded by Danilewsky (1889) ia Southern Russia from 
Upupa epops. 

Habitat. — Blood of Upupa epops orientalis Stuart Baker : 
Portuguese India, Daman. 

166. Hsemoproteus wenyoni de Mello, de Sa, de Sousa, Dias, 
& Noronha. (Fig. 116.) 

"fHxtnoproteus wenyoni, de Mello, de Sa, Sousa, Dias, & Noronha, 
1917,^p. 11-12, pi. i, figs. 18-44 ; 1937 a, p. 100. 

Female gametocyte ovoid or halter-shaped, not displacing 
the nucleus of the infected red blood-corpuscle, stained deep 
blue mth Leishman's stain ; nucleus small, spherical, compact, 
and almost always situated near one pole of the gametocyte 
at its convex border ; pigment granules brown, scattered 
a,ll over the body, rarely confined to one pole. Male gameto- 
cyte stained blue ; nucleus usually central, polymorphic. 

A B C 

Pig. 116. — Hsemoproteus wenyoni de Mello, de Sa, de Sousa, Dias, 
& Noronha. A, female gametocjrte ; B, male gametocyte ; 
C, extra-corpuscular schizont. (After de Mello and others.) 

being a compact chromatic granule or comma- shaped, linear 
or doubled up, or pjrriform, etc. ; pigment yellowish-brown, 
darker than in the female gametocyte, and nearly always 
situated at both the poles. 

Schizogony observed in smears from the liver and the lung, 
but never in the peripheral blood. Schizonts small or of 
medium size. "Acton body " rounded, showing nuclear multi- 
plication. Merozoites not seen. 

Remarks. — The form of the gametocytes, and the occurrence 
of schizogony in smears from the Hver and the lung, make it 
reasonably certain that the parasite is a Hsemoproteus. The 
so-called "Acton body " is probably a detached endothehal 
cell of the blood-vessel. 

Habitat. — Blood and smears from the liver and the lung of 
the tailor-bird, Orthotomus sutorius (Pennant) : Portuguese 
India, Nova Goa. 


167. Hsemoproteus sp. 

■fHsemoproteus (?) sp., Scott, 1926, p. 237, Wenyon, 1926, p. 1365. 
Hsemoproteus (?) sp., Coatney, 1936, p. 90 ; de Mello, 1937 a, 
p. 99. 

Habitat. — Blood of the Sarus crane, Antigone antigone Linn., 
from India, in the Zoological Gardens, London. 

168. Hsemoproteus sp. 

■\Heemoproteus sp., Scott, 1926, p. 237 ; Wenyon, 1926, p. 1368. 
Hsemoproteus sp., Coatney, 1936, p. 91 ; de Mello, 1937 a, p. 99. 

Habitat. — Blood of the Nicobar pigeon, Calosnas nicobarica 
Linn., from India, in the Zoological Gardens, London. 

169. Haemoproteus sp. 

■f Hsemoproteus sp., Donovan (first recorded in Wenyon, 1926, 
p. 1369). 
Hsemoproteus sp., Coatney, 1936, p. 91 ; de Mello, 1937 a, p. 99. 

Habitat. — Blood of the cuckoo, Centropus sinensis (Stephen) : 
India (locality not cited). 

170. HsBmoproteus sp. 

■f Hsemoproteus sp., Scott, 1926, p. 237 ; Wenyon, 1926, p. 1369. 
Hsemoproteus sp., Coatney, 1936, p. 91 ; de Mello, 1937 a, p. 99. 

Habitat. — Blood of the golden-headed chloropsis, Chloropsis 
aurifrons (Temm.), from India, in the Zoological Gardens, 

171. Hsemoproteus sp. 

■\ Hsemoproteus sp., Castellani & Willey, 1905, p. 385, pi. xxiv, fig. 4, 
Hsemoproteus sp., Wenyon, 1926, p. 1371. 

Rare parasites of an elongate shape, with two or three 
vacuoles, free in the plasma. Sometimes the free parasite 
appears to be surrounded by a clear halo. Nucleus oval, 

Remarks. — The parasite appeared to be non-motile, but 
showed a slight change of shape. 

Habitat. — Blood of the common babbler, Turdoides striatus 
striatus (Swainson) : Ceylon. 

172. Hsemoproteus sp. 

'\ Hsemoproteus sp., Plimmer, 1916, p. 85. 
Hsemoproteus sp., Wenyon, 1926, p. 1371 ; Coatney, 1936, p. 93. 

Habitat. — Blood of yellow- fronted barbet, Cyanops flavifrons 
(Cuvier), from Ceylon, in the Zoological Gardens, London. 


173. Hsemoproteus sp. 

'\Hsemoproteus sp., Pliinnier, 1913, p. 148. 
Hsemoproteus sp., Wenyon, 1926, p. 1371 ; Coatney, 1936, p. 93. 

Habitat. — Blood of the wandering tree-pie, Dendrocitta 
rufa vagabunda Latham, from India, in the Zoological Gardens, 

174. Hsemoproteus sp. 

■\Hsemoproteus sp., Scott, 1926, p. 237 ; Wenyon, 1926, p. 1375. 
Hsemoproteus sp., Coatney, 1936, p. 95 ; de Mello, 1937 a, p. 99. 

Habitat. — Blood of Pallas's fishing eagle, Haliaetus leucory- 
pjius Pallas, from India, in the Zoological Gardens, London. 

175. Hsemoproteus sp. 

■\Hsemoproteus sp., Donovan (first recorded in Wenyon, 1926, 
p. 1388). 

Habitat. — Blood of Hemidactylus sp. : India (locaHty not 

176. Hsemoproteus (?) sp. 

■f Hsemoproteus (?) sp., Scott, 1926, p. 237 ; Wenyon, 1926, p. 1377^ 

Habitat. — Blood of the Ceylon loriquet, Coryllis beryllinus 
(Forster), from Ceylon, in the Zoological Gardens, London. 

177. Hsemoproteus sp. 

■^Hsemoproteus sp., Plimmer, 1913, p. 148 ; Knowles, 1928, p. 372. 

Habitat. — Blood of the red-breasted parrakeet, Psittacula 
alexandri fasciatus (Miiller), from India, in the Zoological 
Gardens, London ; also from parrots in the Ahpore Zoological 
Gardens : Bengal, Calcutta. 

178. Hsemoproteus sp. 

^Hsemoproteus sp., de Mello, 1935, p. 470 ; 1937 a, p. 100. 

Female gametocyte stained Hght prussian-blue with 
Leishman's stain ; nucleus pale rose. Male gametocyte 
stained Ught bluish-grey or colourless ; nucleus a little larger 
than in the female ; pigment coffee-coloured, in granules, 
seldom in rods, scattered singly or in clusters. Shows the peculi- 
arity of almost completely hsemolysing the infected cell. 

Remarks. — Hsemoproteids have been previously recorded 
from several species of Strix from dijfferent parts of the world, 
and in view of the possibihty that it might be specifically 


identical with one or the other of those parasites, de Mello 
has not given a name to the form observed by him. 

Habitat — Blood of Strix ocellata (Lesson) : Portuguese 
India, Pragana. 

179. Hsemoproteus sp. 

■\Hsemoproteus sp., Knowles, 1928, p. 372. 

Habitat. — ^Blood of water-fowls, buzzards, Java sparrows, 
munias, hawks, parrakeets, and canaries in the Ahpore 
Zoological Gardens : Bengal, Calcutta. 

180. Hsemoproteus (?) SBgithinae de Mello. 

•f Hsemoproteus segitMnse, de Mello, 1935 6, p. 471 ; 1937 a, p. 100. 

Female gametocyte stained Hght blue with Leishman's 
stain ; nucleus ovoid, central or subcentral ; pigment in 
minute granules, scattered over the body. Male gametocjrfce 
colourless ; nucleus oval, subcentral or central ; pigment in 
fine granules, dust-hke, collected mostly at the poles. Red 
cell hypertrophied, with its nucleus displaced. 

Habitat. — Blood of jEgithina tiphia (Linn.) : Portuguese 
India, Nagoa (Salcete). 

181. Haemoproteus (?) antM de Mello. 

"^Hsemoproteus anthi, de Mello, 1935 &, p. 474 ; 1937 a, p. 100. 

Female gametocyte stained hght blue, but not uniformly, 
with Leishman's stain ; nucleus compact, central or sub- 
central, stained deep rose. Male gametocyte not stained ; 
nucleus oval or triangular, central. Pigment in both kinds 
of gametocytes brown, coffee-coloured, irregularly scattered, 
seldom located at the poles. Infected red cell hypertrophied, 
with its nucleus displaced. 

Remarks. — De Mello thinks that the parasite is perhaps 
similar to the unnamed Hsemoproteiis recorded from Anthtis 
trivialis by Galli Valerio (1902) in Europe, from A. japonicus 
by Ogawa (1911) in Japan, and from A. pratensis by Nieschulz 
(1921, 1922) in Heligoland. 

Habitat. — Blood of Anthus richardi rufulus Vieillot : Portu- 
guese India, Nagoa (Salcete). 

182. Hsemoproteus (?) bramse de Mello. 

■fHsemoproteus sp., Donovan (first recorded in Wenyon, 1926, 

p. 1366). 
■^Hsemoproteus bramse, de Mello, 1935 h, pp. 474-5 ; 1937 a, p. 100. 

Female gametocyte halter-shaped, slender, more or less 
irregular, somewhat constricted in the middle, rarefy oval. 


cystoplasm homogeneous, stained blue with Leishman's stain, 
more deeply at the poles ; nucleus oval, central or subcentral, 
stained pale rose ; pigment granules isolated or in clusters, 
irregularly scattered. Male gametocyte halter-shaped, comma- 
shaped, or oval ; nucleus without definite outline, often 
scarcely visible ; pigment granules fewer than in the female 
gametocyte, generally at the poles, but sometimes over the 
whole body. Infected red cell unaltered or sHghtly hyper- 
trophied, with its nucleus displaced. 

Habitat. — Blood of Athene hrama Temm. : India (locahty 
not cited) ; Portuguese India, Nagoa (Salcete). 

183. Hsemoproteus (?) centropi de Mello. 

-\Hsemoproteus centropi, de Mello, 1935 b, p. 471 ; 1937 a, p. 100. 

Female gametocyte with alveolar cytoplasm, stained light 
blue with Leishman's stain ; nucleus central, pale rose ; 
pigment yellow-brown, in large or minute dust-like granules, 
situated either at the poles or irregularly scattered over the 
body. Male gametocyte colourless, or with a sHghtly pale 
blue tinge in the periphery or at the poles ; nucleus central, 
much larger than in the female gametocyte, and containing 
minute distinct chromatic dots or an irregular spirematic 
thread ; pigment very irregularly distributed, but showing 
a tendency for a polar location. Infected red cell hypertrophied, 
with its nucleus displaced. 

Remarks. — Donovan recorded an unnamed Hsemoproteid 
from Centropiis sinensis from India, and unnamed Hsemopro- 
teids have also been recorded from two other species of the 
same genus from other parts of the world. 

Habitat. — Blood of Centropus sinensis parroti Stresemann : 
Portuguese India, Diu. 

184. Hsemoproteus (?) dicruri de Mello. 

■fHsemoproteus dicruri, de Mello, 1935 6, p. 473 ; 1937 a, p. 100. 

Female gametocyte stained bluish with Leishman's stain, 
but not uniformly, the poles and the convex border remaining 
unstained ; nucleus small, subcentral, stained pale rose ; 
pigment yeUow-brown, in granules or rods, collected at the 
poles. Male gametocyte ovoid, not stained ; nucleus rod-like, 
subcentral, sometimes indistinct, stained pale rose ; pigment 
in large granules, scattered or collected in clusters at the 
poles. Infected red cell hypertrophied, with its nucleus 

Habitat. — Blood of Dicrurus macrocercus macrocercus Vieil- 
lot : Portuguese India, Pragana. 


185. Hsemoproteus (?) elani de Mello. 

^Hsemoproteus elani, de Mello, 1935 b, p. 472 ; 1937 a, p. 100. 

Female gametocyte stained uniformly blue with Leishman's 
stain ; nucleus compact, small, ovoid, rarely triangular, sub- 
central, stained reddish. Male gametocyte not stained ; 
nucleus large, granular, without a definite outline. Pigment 
of sepia colour, disposed in both male and female gametocytes 
in a characteristic manner, being in the form of large mulberry- 
like masses at the poles. Infected red cell hypertrophied, 
with its nucleus displaced. 

Habitat. — Blood of Elanus casruleus vociferus Latham : 
Portuguese India, Daman. 

186. Hsemoproteus (?) gallinulse de Mello. 

■\H3em0pr0teus gallinulse, de Mello, 1935 h, p. 469 ; 1937 a, p. 100. 

Female gametocyte with vacuolated cytoplasm, stained 
deep blue with Leishman's stain ; nucleus stained pale rose ; 
pigment brownish-black, in granules of different sizes scattered 
all over the body, rarely collected at the poles, sometimes 
grouped in clusters. Male gametocyte stained Kght blue, 
rather colourless, roundish or oval, almost never halter-like ; 
pigment in irregular clusters collected at the poles. Infected 
red cells hypertrophied, with the nucleus displaced. No 
schizogonic forms observed in lung-smears. 

Habitat. — Blood of Gallinula chloropus (Linn.) : Portu- 
guese India, lake of CaromboHm. 

187. Hsemoproteus (?) halcyonis de Mello. 

1f Hsemoproteus halcyonis, de Mello, 1935 b, p. 474 ; 1937 a, p. 100. 

Female gametocyte halter-like, oval, or cordiform, vacuolated 
or not, stained deep blue with Leishman's stain ; nucleus oval, 
central or subcentral ; pigment at the poles, sometimes in 
the centre or near the periphery. Male gametocyte not 
stained ; nucleus indistinct, oval or comma-shaped, central 
or subcentral ; pigment scattered over the body, often denser 
at the periphery. Infected red cell shghtly hypertrophied, 
sometimes not at all, with its nucleus displaced. 

Habitat. — Blood of Halcyon smyrnensis (Linn.) : Portu- 
guese India, Canacona. 

188. Hsemoproteus (?) halcyonis var. fuscse de Mello & da 

Fonseca. (Fig. 117.) 

'^Hsemoproteus halcyonis juscse, de Mello & da Fonseca, 1937, 
pp. 215-16, text-figs. 

Female gametocyte oval, fusiform or like a slender halter 
when young, definitely halter-like when full grown ; often 



the halteridia are somewhat irregular, and when fully grown 
both poles of the gametocji^e fuse together, surrounding the 
nucleus of the red cell ; stained definitely blue with Romanow- 
sky's stain ; nucleus oval, generally central ; pigment granules 
blackish-brown or coffee-brown, with irregular distribution. 
Male gametocyte oval when young, halter-like when full- 
grown, sometimes completely surrounding the nucleus of 
the red cell, not stained or slightly blue with Romanowsky's 

A B C D 

Fig. 117. — Hsenioproteus (?) halcyonis -var. fuscse de Mello & da Fonseca. 
A, B, female gametocytes ; C, D, male gametoeytes. 
(After de Mello & da Fonseca.) 

stain ; nucleus large, central or subcentral, stained rose by 
Leishman's stain, reddish by May-Grunwald Giemsa stain ; 
pigment granules variable in size and location, showing 
a tendency to be callected at the poles. Infected red cell 
not altered when the parasite is young ; when the latter is 
full grown the nucleus of the red cell is displaced to the 

Habitat. — Blood of Halycyon smyrnensis fusca (Bodd.) : 
Portuguese India, Santo Estevam. 

189. Heemoproteus (?) lanii de Mello. (Fig. 118.) 

Hsemoproteus lanii, de Mello, 1937 a, pp. 102-3, pi. i, fig. 2. 
Female gametocyte halter-shaped, staining blue with 

A B 

Fig. 118. — Hsemoproteus (?) lanii de Mello. A, female gametocyte; 
B, male gametocyte. (After de Mello.) 


Leishman's stain ; nucleus small, oval or triangular, generally 
subcentral, not staining or very lightly with Leishman's 
stain, staining well with May-Grunwald Giemsa stain ; pig- 
ment granules large, scattered irregularly over the body. 
Male gametocyte irregularly halter-shaped, almost quadrangu- 
lar, not stained or very light blue ; nucleus large, central, 
without definite outline, with chromatinic masses irregularly 
scattered or disposed in a reticulum ; pigment granules very 
minute, located at the poles. Infected red cell hypertrophied, 
with its nucleus displaced. 

Bemarks. — Unnamed species of Hasmoproteus have been 
previously recorded from many different species of the genus 
Lanius in different parts of the world. 

Habitat. — Blood of Lanius schach erythronotus Vigors : Por- 
tuguese India, Ponda. 

190. Hsemoproteus (?) otocompsse de Mello. 

"fHsemoproteus otocompsse, de Mello, 1935 b, p. 473 ; 1937 a, p. 100. 

Female gametocyte with cytoplasm stained pale blue with 
Leishman's stain and rose with May-Grunwald Giemsa stain, 
seldom vacuolated ; nucleus spherical, subcentral, seldom 
elongated, situated on the convex border ; pigment scattered 
over the body. Male gametocyte not stained with Leishman's 
stain, pale rose with May-Grunwald Giemsa stain ; nucleus 
very large, granular, without definite outhne, subcentral ; 
pigment at the poles. Infected red cell hypertrophied, with 
its nucleus displaced. 

Bemarks. — According to de Mello the parasite is perhaps 
identical with the unnamed Hsemoproteus of Picnonotus harhatus 
recorded by A. and M. Leger in Senegal. 

Habitat. — Blood of Elathea jocosa (Linn.) [^Otocompsa 
emeria (Linn.)] : Portuguese India, Malim (Bardez). 

191. Haemoproteus (?) pastoris de Mello. 

■\Heemoproteus vastoris, de Mello, 1935 b, p. 470. 
Hsemoproteus sp., Coatney, 1936, p. 98. 
Heem,oproteus pastoris, de Mello, 1937 a, p. 100. 

Female gametocyte stained blue with Leishman's stain, 
with a small ovoid nucleus, central or subcentral, stained 
pale rose ; halter-shaped or ovoid, and in the latter case 
nucleus of the host- cell completely displaced ; pigment 
scattered all over the body. Male gametocyte colourless or 
very shghtly blue, halter-shaped, ovoid or irregularly deformed ; 
nucleus indistinct, composed of chromatic masses disposed 
like the spokes of a wheel, or in irregularly weaved threads ; 
pigment polar in halter-shaped forms, scattered over the 


body in ovoid or deformed gametocytes. Infected red cell 
hypertophied, with its nucleus displaced. 

Habitat. — Blood of Pastor roseus (Linn.) : Pokttjguese 
India, Pragana. 

192. HsBmoproteus (?) platalesB de Mello. 

■\Hsemoproteus platalese, de Mello, 1935 6, p. 471. 
Hsemoproteus sp., Coatney, 1936, p. 99. 
Hsemoproteus platalese, de Mello, 1937 a, p. 100. 

Female gametocyte with alveolar cytoplasm, stained deep 
blue with a greenish tone with Leishman's stain ; nucleus 
central ; pigment coffee-oHve tone, scattered over the body, 
showing a tendency to undergo solution and to diffuse through 
the cytoplasm. Male gametocyte not stained, or very sHghtly 
greyish-blue, the margins at the poles being often the only 
stained part ; pigment in minute granules, often in clusters, 
generally at the poles. Infected red cell hypertrophied, 
with its nucleus displaced. 

Habitat. — Blood of Platalea leucorodia major Temm. & 
Schleg. : Poktttgijese India, Diu. 

193. Hsemoproteus (?) tephrodornis de Mello. 

■fHsemoproteus tephrodornis, de Mello, 1935 b, p. 473 ; 1937 a, p. 100. 

Female gametocyte halter-shaped or ovoid, stained blue 
with Leishman's stain, but not uniformly ; nucleus oval, 
central or subcentral. Male gametocyte oval, stained Hght 
greyish ; nucleus small, indistinct. Pigment black or sepia 
coloured, in granules or rods disposed in lines or clusters 
along the concave border. Infected red cell hypertrophied, 
with its nucleus displaced. 

Habitat. — Blood of Tephrodornis pondicerianus pondicerianus 
(GmeHn) : Portttguese India, Pragana. 

194. Hsemoproteus (?) thereicerycis de MeUo. 

■f Hsemoproteus thereicerycis, de Mello, 1935 b, p. 470 ; 1937 a, p. 100. 

Female gametocyte halter-shaped, ovoid or spherical, 
cytoplasm finely vacuolated, stained blue with Leishman's 
stain ; nucleus hardly distinguishable with Leishman, well 
stained with May-Grunwald Giemsa stain, spherical, central or 
subcentral, lodged in a definite vacuole ; pigment granules of 
different sizes scattered over the body. Male gametocyte 
always spherical, colourless with Leishman's stain, rose with 
May-Grunwald Giemsa stain ; nucleus very small, comma- 
shaped ; pigment granules at the poles a little larger than those 
scattered over the body. Infected red cell hypertrophied, with 


its nucleus displaced. Shows a marked tendency to hsemolyse 
the infected cell and become free, taking in this free condition 
aberrant forms like deformed halteridia or spheres. 

Habitat. — Blood of Thereiceryx zeylanicus inornata Walden : 
PoRTTJGTJBSE India, Corlim (Ilhas). 

195. HsBmoproteus (?) thereieerycis var. zeyloniea de Mello. 

•fHeemoproteus thereieerycis var. zeyloniea, de Mello, 1935, p. 470 ; 
1937 a, pp. 100, 101-2. 

Female gametocyte with vacuolated cytoplasm, stained 
dark blue with Leishman's stain ; nucleus round, seldom 
elongated, central or subcentral ; pigment granules scattered, 
minute, often in clusters. Male gametocyte colourless ; 
nucleus large and without definite outline, always central, 
and containing chromatic masses with irregular disposition ; 
pigment granules very minute, situated at the poles. 
Infected red cell hypertrophied, with its nucleus displaced. 

Remarks. — This variety differs from the preceding species as 
regards the structure of the male gametocyte, and does not 
show the same tendency to hsemolyse the host- cell. 

Habitat. — ^loodoi Thereiceryx zeylanicus zeylanicus (Gmelin) : 
Portuguese India, Malim (Bardez) ; blood of Thereiceryx 
viridis Bodd. : Portuguese India, Ponda. 

Genus LEUCOCYTOZOON Danilewsky, 1889. 

Leucocytozoon, Danilewsky, 1889, p. 23 ; Berestneff, 1903, pp. 376- 
86 ; Wenyon, 1910, pp. 63-72 ; Franca, 1912, pp. 173-6 ; Mathis 
& Leger, 1912, pp. 77-82 ; Minchin, 1912, pp. 357, 369-71, 
390, 392 ; Moldovan, 1914, pp. 249-62 ; Wenyon, 1926, pp. 903-8; 
Kudo, 1931, p. 288 ; Calkins, 1933, p. 566 ; de Mello, 1935 a, 
pp. 355-7 ; 1935 b, pp. 68-73 ; Reichenow, 1935, p. 375. 

Schizogony takes place in the internal organs of the host, 
probably in the endothelial cells of the blood-vessels (as in 
Heemoproteus). Certain cells in the peripheral blood, which 
were originally thought to be leucocytes (hence the name of 
the genus) but are now believed to be young erythrocytes, 
in which the colouring matter has not yet developed, contain 
the gametocytes in various stages of growth. No pigment 
is produced. Micro- and macrogametocytes are differentiated, 
the nucleus of the microgametocyte is large and contains 
numerous diffuse chromatin granules, while that of the macro- 
gametocyte is compact and contains a definite karyosome. 
The host-cell is profoundly modified into an elongate fusiform 
body, much larger than the normal red blood- corpuscle, and 
contains the elongate and hypertrophied nucleus of the cell 
and the elongate gametocyte. The gametocytes leave the 


host-cell, the microgametocyte producing the microgametes 
by a process of flagellation and the macrogametocyte rounding 
o£F to form a macrogamete. Finally, a large unpigmented 
ookinete is formed. Blood parasites of various birds. 

Remarks. — ^Wenyon (1926) gives a useful summary of the 
previous Hterature on this genus and places it in the same 
family with Heemoproteus, from which it differs chiefly in the 
absence of pigment in the gametocytes and the peculiar modi- 
fication of the host-cell. Reichenow (1931), stressing the 
affinities with the Coccidia, places it in a family by itself 
under Eimeridba. I have followed Wenyon, Kudo, and 
Calkins in placing it here. De Mello (1937 a) has surveyed 
the previous literature deahng with the genus, and come 
to the conclusion that species with a rounded form should be 
placed in an independent genus from Leucocytozoon, which 
should be restricted to the fusiform species. 

196. Leucocytozoon ardeolse de Mello. 

'\ Leucocytozoon ardeolse, de Mello, 1937 a, p. 106. 

Form round. No sexual differentiation made out. Cyto- 
plasm with very minute alveoli, staining deep blue with 
Leishman's stain ; nucleus circular, staining pale rose. 

Habitat. — Blood of Ardeola grayi Sykes : Portuguese 

197. Leucocytozoon chloropsidis de Mello. (Fig. 119.) 

\ Leucocytozoon chloropsidis, de Mello, 1935 a, pp. 356-7, pi. xliii, 
fig. 2 ; 1937 a, p. 105. 

Form round or oval. Cytoplasm alveolar. Sexual dimor- 
phism of the gametocytes not shown by the cytoplasm, which 
stains deep blue in both the male and female, but by the 

A B 

Fig. 119. — Leucocytozoon chloropsidis de Mello. 
A, female gametocyte ; B, male gametocyte. (After de Mello.) 

appearance of the nucleus. In the male gametocytes the 
nucleus is long, thread-like, irregular, and stains violet by 
Eiomanowsky stains. In the female gametocytes the nucleus 



is roundish or oval, weakly stained pale rose or not stained 
at all. Infection very scanty. 

Habitat. — Blood of the golden-headed chloropsis, Ghloropsis 
aurifrons davidsoni Stuart Baker : Poktugtjbse India, 

198. Leucocytozoon coraeiaB de Mello & Afonso. (Fig. 120.) 

■fLeucocytozoon coraciae benghalensis, de Mello & Afonso, 1935 b, 
pp. 71-2, pi. ii ; de Mello, 1937 a, p. 105. 

Form of the adult gametocyte ovoid, included in a fusiform 
cell. Cytoplasm finely alveolar stains blue with Romanowsky 
stains, but lighter than in i^. melloi. Ratio of length to breadth is 
5 to 2 in intracellular forms and 7 to 4 in free forms. Nucleus 

A B 

Fig. 120. — Leucocytozoon coraciee de Mello & Afonso. 

A, female gametocyte ; B, male gainetocyte. 
(After de Mello and Afonso.) 

ovoid or bean-shaped, central or subcentral. Sexual dimor- 
phism is recognizable in the structure of the nucleus as stained 
by iron hsematoxyhn. The nucleus of the female gametocyte 
is small, with a karyosome and a centriole or only with centriole. 
The nucleus of the male gametocyte is much larger, often oval, 
elongated, or even bean-shaped, filled with granules, but without 
a centriole. 

Remarks. — No young forms or schizogony stages were 
recognized. Specimens of this species were far more abundant 
(in the ratio of 7 to 1) than those of L. melloi described from 
the same host. 

The poles of the fusiform host-cell are stained greyish-blue 
by Romanowsky stains, and stained Hghtly by Heidenhain's iron 


hsematoxylin. The nucleus of the host-cell is attached to one 
side of the parasite, giving it a bean- shaped form. The 
parasite is really ovoid, as can be seen in free forms or where 
the nucelus of the host-cell is still central. In many cases 
one or both poles of the host-cell are lost, and it appears to 
be represented only by its nucleus. 

Habitat. — Blood or smears from the lungs of Coracias 
benghalensis benghnlensis (Linn.) : Portuguese India, Corhm. 

199. Leucocytozoon enriquesi de Mello. 

jLeucocytozoon enriquesi, de Mello, 1937 a, p. 106. 

Form round. Differs from the preceding species in the 
nucleus being roundish, ovoid or sausage- shaped and always 
well stained in the female gametocyte ; and in the nucleus 
being very large, circular, and occupying the greater part of 
the bod}'", and stained pale rose, dust-like and with a more 
deeply staining body, in the male gametocyte. 

Habitat. — Blood of Ghloropsis jerdoni Blyth : Portuguese 

200. Leucocytozoon (?) melloi, sp. nov. (Fig. 121.) 

•\Leucocytozoon (?) sp., "type B," de Mello & Afonso, 1935 b, 
pp. 72-3, pi. ii ; de Mello, 1937 a, p. 105. 

Form of the adult gametocyte spherical, attached to the 
nuclear substance of the host- cell, the rest of the cell not to be 
seen. Cytoplasm compact, stained deep blue by Romanowsky 
stains, and much darker than L. coracise by iron haematoxylin ; 

Fig. 121. — Leucocytozoon (?) melloi, sp. nov. 

A, B, forms with nucleus of the host-cell attached ; G, free form. 
(After de Mello and Afonso.) 

containing small or large vacuoles, which do not take the 
stain. Only one type of gametocyte recognizable, with 
a nucleus containing a karyosome. 

Remarks. — Forms are of different sizes, the free ones being 
absolutely spherical, and not ovoid as in L. coracise from the 
same host. The relationship of the nucleus of the host- cell 



to the parasite is entirely similar to that of the host- cell and 
the previous species. 

Marcel Leger (1913) and Laverallan and Marullaz (1914) 
have previously described Leucocytozoons of a rounded form, 
and interpreted them as different stages of the typical form. 
De Mello and Afonso (1935 b) are, however, of the opinion that 
they are definitely deaHng with two independent species, 
of which the first one may be classified as a Leucocytozoon, 
while the second has enough characters to constitute an 
independent genus, to which the name of Marcel Leger should 
be attached. As they have not given a definite name to the ' 
species, but have simply described it as " type B," I have 
for the present kept it in the genus Leucocytozoon and have 
given the name melloi to the species. 

Habitat. — Blood and smears from the lungs of Coracias 
benghalensis benghalensis (Linn.) : Portuguese India, CorHm. 

201. Leucocytozoon molpastis de Mello. 

■^Leucocytozoon molpastis, de Mello, 1937 a, pp. 106-7, pi., fig. 1. 

Form round. Cytoplasm with very few minute vacuoles, 
not showing sexual differentiation. Nucleus circular, with 
a well-marked membrane and a conspicuous central granule 
in the female gametocyte ; oval, with chromatin disposed in 
variously arranged threads in the male gametocyte. 

Habitat. — Blood of Molpastes cafer cafer (Linn.) : Portu- 
guese India, Ponda. 

202. Leucocytozoon sp. 

"fLeucocytozoon sp., Donovan (first recorded in Wenyon, 1926, 
p. 1366). 
Leucocytozoon sp., de Mello, 1937 a, p. 105. 

Habitat. — Blood of the Indian little owl, Athene brama 
(Temm.) : India (locahty not cited). 

203. Leucocytozoon sp. 

■f Leucocytozoon sp., Donovan (first recorded in Wenyon, 1926, 
p. i373). 
Leucocytozoon sp., de Mello, 1937 a, p. 105. 

Habitat. — Blood of the falcon, Falco sp. : India (locahty 
not cited). 

204. Leucocytozoon sp. 

■f Leucocytozoon sp., Scott, 1926, p. 237 ; Wenyon, 1926, p. 1375. 
Leucocytozoon sp., de Mello, 1937 a, p. 105. 

Habitat. — Blood of the rufous-chinned laughing-thrush, 
lanthocincla rufogularis Gould, from India, in the Zoological 
Gardens, London. 


205. Leucocytozoon sp. 

■\Leucocytozoon sp., Knowles, 1925. 

Leucocytozoon sp., Wenyon, 1926, p. 1377 ; de Mello, 1937 a, 
p. 105. 

Habitat. — Blood of the babbler, Leiothrix lutea (Scop.) : 
Bengal, Calcutta. 

206. Leucocytozoon sp. 

■f Leucocytozoon sp., Plimmer, 1914, p. 190. 
Leucocytozoon sp., Wenyon, 1926, p. 1379 ; de Mello, 1937 a, p. 105. 

Habitat. — Blood of the chat, Oreicola ferrea (Gray), from 
India, in the Zoological Gardens, London. 

207. Leucocytozoon sp. 

"[Leucocytozoon sp., Plimmer, 1917, p. 32. 
Leucocytozoon sp., Wenyon, 1926, p. 1382 ; de Mello, 1937 a, p. 105. 

Habitat. — Blood of the finch, Propasser rhodochrous (Vigors), 
from India, in the Zoological Gardens, London. 

208. Leucocytozoon sp. 

"l Leucocytozoon sp., Knowles, 1928, pp. 379, 381. 

Remarks. — The host-cells, although very much enlarged 
and showing the characterististic squeezing-out of the nucleus 
by the parasite against the cell-membrane, did not show the 
drawn-out tapering ends. Knowles thinks this may have 
been due to post-mortem changes, as the material was not 

Habitat. — Blood from the heart and other viscera of Peking 
robins in the Ahpore Zoological Gardens : Bengal, Calcutta. 

209. Leucocytozoon (?) sp. 

■fLeucocytozoon (?) sp., de Mello, 1937 a, p. 105. 

Form round. Female gametocyte with very small vacuoles,^ 
and staining deep blue with Leishman's stain; nucleus ovoid, 
granular, staining rose colour. Male gametocyte with large 
vacuoles, and staining Hght blue ; nucleus in the form of 
an irregular thread. 

Remarks. — According to de Mello, the form is perhaps 
similar to L. annelloise Cleland, 1912, described from Oriolus 
Sagittarius in Austraha. 

Habitat. — Blood of Oriolus oriolus kundoo Sykes : Poetf- 
GUESE India. 



210. Leucocytozoon (?) sp. 

■fLeucocytozoon (?) sp., de Mello, 1937 a, pp. 105-6, 

Form round. No sexual differentiation. Great tendency 
to aberrant forms in addition to round ones of regular t3rpe. 
Nucleus crescentic or irregular and very large in some speci- 
mens ; oval, with a more deeply staining line or point in others. 

Habitat. — Blood of Oriolus xanthomus xanthornus (Linn.) : 
Portuguese India. 

2. Family PLASMODIID^ Mesnil, 1903. 

Schizogony takes place within the red blood- corpuscles of 
s, Vertebrate host. During the growth of the trophozoite, 
pigment, known as hsemozoin, is formed from the haemoglobin 
of the corpuscle. Gametocytes also occur in the red blood- 
curpuscles and contain pigment. Further development takes 
place in the body of a mosquito. Microgametes are produced 
by exflagellation. Zygote becomes a motile ookinete, and 
later encysts as an oocyst. The oocyst grows enormously 
in size, and innumerable sporozoites are produced, without 
the formation of sporoblasts or sporocysts. 

The family, according to many authorities, contains a single 
genus, Plasmodium, as they consider Plasmodium Marchiafava 
& Celh, Laverania Grassi & Feletti, and Proteosoma Labbe as 
congeneric. Doflein (1916) and Reichenow (1929) consider 
these as distinct, and Reichenow also includes the genus 
Dactylosoma in the family. According to Reichenow it would 
be desirable to retain the generic name Proteosoma for the 
malarial parasites of birds, and to refer the malarial parasites 
of the reptiles also to it. He also considers it justifiable to 
place the human tropical parasite of mahgnant tertian malaria 
in a separate genus, Laverania. Both the genera Proteosoma 
and Laverania show some resemblance to Coccidia in that the 
gametocytes are spherical, oval, vermiform or sausage-shaped, 
whilst in Plasmodium the gametocytes are circular and disc- 
like. The genus Laverania, which includes parasites of man 
and anthropoid apes, is not easily marked off from the 
Proteosoma of birds. Slender gametocytes, like those charac- 
teristic of Laverania, occur also in some species of Proteosoma, 
and the characteristic that schizogony takes place in the internal 
organs in the case of Laverania has been found to occur, 
according to the recent researches of Hartmann, in Proteosoma 

Of the English authors, Thomson and Woodcock (1922) 
justified the retention of Laverania as a separate genus on the 
ground that the gametocyte or crescent is enclosed by a capsule, 


the male crescent possessing a more delicate capsule than 
the female crescent. Wenyon (1926) is not convinced that 
any such capsule exists, and is of the opinion that the Hfe- 
cycles of the three species of malarial parasites are so similar 
in every respect that, even if a capsule did exist, it would 
not justify generic distinction. In my opinion the life-cycle 
is bound to be similar within the same family, and the difference 
in form of the gametocyte is a sufficiently important feature 
to serve as a basis for generic distinction. Stiles (1928) 
has announced that the International Commission of Zoological 
Nomenclature has placed both Laverania and Plasmodium 
in the official list of generic names. 

History of the Discovery of Malarial Parasites and the part 
played by the Mosquitoes. — It will not be out of place to 
briefly review here the history of the discovery of malarial 
parasites and of the part played by the mosquitoes in their 
transmission. The study of the HiEMOSPOBiDiA (or Hsemo- 
cytozoa or Hsematozoa as they were then generally called) 
began with the discovery by Ray Lankester in 1871 ofDrepani- 
dium ranarum in the blood of the frog. Human malarial 
parasites are said to have been seen, but their significance was 
not comprehended until Laveran pubHshed his investigations 
(1880). Golgi (1885) demonstrated the relationship existing 
between the life- cycle of the parasites within the human body 
and the occurence of the febrile attack. In India Vandyke 
Carter (1888), Evans (1888), Hehir (1893), Crombie (1894), and 
Ronald Ross (1895) were the earliest to study the malarial 
parasites in man. Ross (1895) observed the process of 
" flagellation " of crescentic parasites in the stomach of 
mosquitoes fed on the blood of a malarial patient. MacCullum 
(1897) found that the " flagella " oi Hsemoproteus (Halteridium) 
and of sestivo- autumnal parasites constitute the male element, 
and serve to impregnate the " pigmented spheres " or female 
element, and further observed in the former that the im- 
pregnated spheres become converted into motile " vermicules." 
Ross (1897), working at Secunderabad, fed mosquitoes upon 
human blood containing " crescents." After examining 
hundreds of mosquitoes fed on malarial blood, with negative 
results, he obtained a few mosquitoes with spotted wings, 
in which he discovered pecuHar pigmented cells lying within 
the walls of their stomachs. The pigment was similar to that 
within the malarial parasites in the blood upon which the 
mosquitoes had been fed. Ross concluded that he had found 
the mosquito which served as a host for the parasite. In 
February 1898 he again referred to his experiments with 
crescentic parasites and dapple- winged mosquitoes. After 
this, working at Calcutta, Ross observed the development 
of the malarial parasite of birds, Proteosoma, in a species of 


Culex (subsequently determined as C. fatigans Wied.), the 
insects being fed on the blood of infected crows, larks, and 
sparrows, and found similar pigmented cells. In July of the 
same year Manson reported to the British Medical Association 
at Edinburgh further observations on behalf of Ross, which 
showed that the encapsulated parasites, on reaching a certain 
size, ruptured and emptied their contents into the coelome, 
and these minute spindle-shaped bodies subsequently accumu- 
lated in the saHvary gland of the insect, thus making it capable 
of transmitting the infection to healthy birds. 

In October 1898 Grassi suspected three species of CuHcidae 
as being carriers of malarial infection, as they were confined 
in their geographical distribution to those regions where malaria 
was prevalent in Italy. It has since been proved that only one 
of these three, viz.. Anopheles claviger, can serve as a host 
for human malarial parasites. A month later Grassi reported 
that Bignami had made an infection experiment with positive 
results. Bignami, Bastianelli, and Grassi (1898) observed 
the development of crescentic parasites in Anopheles claviger, 
and reported that the appearances correspond to those 
described by Ross for Proteosoma on the fourth day in Culex, 
and further that they had successfully infected a person 
with tertian fever by means of infected A. claviger. A few 
weeks later they followed the development of crescentic 
parasites in A. claviger to the formation of sporozoites, their 
escape into the coelome, and their accimiulation in the 
salivary glands of the insect. Koch (1899) reported the 
results of the German Malaria Commission and confirmed 
the development of Proteosoma in Culex as previously described 
by Ross. Daniels (1899) also reported to the Royal Society 
that he had been able to confirm Ross's observations with 

Grassi, Bignami, and BastianelH (1899) observed the 
development of quartan parasites in A . claviger, and Bastianelli 
and Bignami (1899) reported further studies upon the develop- 
ment of tertian parasites in A. claviger, and later extended 
their studies of development in certain other species of 
Anopheles. In September 1899 Bastianelh and Bignami gave a 
detailed description of benign and malignant tertian parasites, 
and their papers were illustrated by the best coloured plates 
published till then, illustrating the development. Ziemann 
(1900) observed the development of the parasites of tropical 
malaria in two species of Anopheles and of tertian parasites 
in one species oi Anopheles, and followed the development up to 
the appearance of sporozoites in the salivary glands of the 
insects. In September 1900 Manson reported a positive 
infection experiment with tertian-fed Anopheles imported 
from Rome, the insects being permitted to bite his son. 
Luhe (1900) showed the correspondence of the life-cycle of the 


malarial parasites with that of Coccidia, and introduced 
a terminology which is in use up till now. Nuttall (1901) 
gave a useful summary of the researches on malaria, recording 
the various works in a chronological order, and the interested 
reader may be referred to it for fuller details. Laveran (1907) 
also reviewed the pubhshed literature. 

Key to Indian Genera. 

1 (4). Schizogony mostly in the red blood- 

corpuscles in* the internal organs ; 
gametocytes spherical, ovoid, vermi- 
form or sausage-shaped 2. 

2 (3). Parasites of birds or reptiles ; gameto- [p. 247. 

cytes spherical, ovoid, or vermiform . . Pkoteosoma Labbe, 

3 (2). Parasites of man and anthropoid apes ; 

gametocytes crescentic or sausage- [Feletti, p. 259. 

shaped Laveeania Grassi & 

4(1). Schizogony takes place in the red blood- 
corpuscles in the peripheral blood ; 
gametocytes circular and disc-like in 

outline. Parasites of man and other [fava & Celli, p. 270. 
naammals Plasmodium Marchia- 

Genus PROTEOSOMA Labbe, 1894. 

PseudovaciMl3e-\-Polymitus-\-Pseudospirilles, Danilewsky, 1889. 
Heemamceba (part), Grassi & Feletti, 1890, p. 463 ; 1891 a, p. 465 ; 

1892, p. 10. 
Hszmoprot&us (part), Kruse, 1890, p. 359. 
Hxmoproteus vars. & & c, Celli & San Felice, 1891, pp. 541-8, 

pis. vii, viii. 
Corps spherique-\- Corps a flag elles-{- Corps en rosette (part), Laveran, 

Cytosporon+Polymitus malaria avium, Danilewsky, 1891, p. 758. 
Proteosoma, Labbe, 1894, pp. 142, 157. 
Heemoproteus, Labbe, 1899, p. 79. 

Hsemoproteus (Proteosoma), Minchin, 1903, pp. 250, 257, 265, 267. 
Plasmodium, Minchin, 1912, p. 358. 

Proteosoma, Doflein, 1916, pp. 896-901 ; Franca, 1917, pp. 221-7. 
Plasmodium, Castellani & Chalmers, 1919, p. 513; Taliaferro, 1925, 

pp. 742-89 ; Wenyon, 1926, pp. 977-81. 
Proteosoma, Calkins, 1926, p. 444. 
Plasmodium, Hartmann, 1927, pp. 1-7 ; Knowles, 1928, pp. 412- 

13, p. 442 ; Hegner, 1929, pp. 59-82 ; Manwell, 1929, pp. 308-45. 
Proteosoma, Reichenow, 1929, pp. 983-7. 
Plasmodium, ManweU, 1930, pp. 381-3; Kudo, 1931, p. 287; Ed. 

& Et. Sergent & Catanei, 1931, pp. 399-437 ; Calkhis, 1933, 

pp. 407, 566 ; Giovannola, 1934, pp. 1-10. 
Proteosoma, Reichenow, 1935, pp. 376-7. 

Schizogony takes place in the red blood-corpuscles, usually 
in the internal organs, but rarely in the peripheral blood. 
The fully developed schizont may more or less displace the 
nucleus of the host-cell or not affect it at all. Gametocytes 
spherical, ovoid or vermiform, in some species closely re- 
sembling those of Laverania. Sporogony as in the human 


malarial parasities of the genus Laverania and Plasmodium, 
Blood-parasities of birds and reptiles. 

Remarks. — Celli and San Felice (1891) reported three kinds 
of malarial parasites in birds, but these were not described 
sufficiently adequately to be recognizable. Till 1927 the 
organisms continued to be considered as belonging to a single 
species, usualty called Plasmodium, precox. Hartmann (1927 6), 
however, beheves that at least three species exist. For one 
of these, isolated from " Whitmore " strain, he retains the 
name P. prsecox ; the second, known as " Hartmann " strain, 
he named P. cathemerium ; and the third strain, obtained by 
Huff in Virginia, was named P. inconstans. Evidence has 
been secured by Manwell (1929) that the " Whitmore " strain 
may consist of two distinct varieties or species. He seems 
to have estabhshed that the " Whitmore " strain of malaria, 
originally isolated by Whitmore from a New York sparrow 
in 1913, is a variety of P. inconstans, and is distinct from 
P. prsecox, which was isolated by Huff in 1926 from canaries 
originally infected T\dth the " Whitmore " strain. Ed. and 
Et. Sergent and Catanei (1929) described another species 
from Algerian birds as P. rouxi, and Huff (1930) yet another 
as P. elongatum from canary and sparrow from the United 
States of America. Manwell (1930) regards P. cathemerium 
Hartmann and P. elongatum Huff as valid species, and 
P. inconstans Hartmann, the " Whitmore " strain, and the 
" German " strain as three strains or varieties of P. prsecox. 
Thus according to him there are three species, namely, P. prse- 
cox, P. cathemerium, and P. elongatum. These three species 
differ from one another in morphology, length of incubation 
period, size of oocysts produced in the mosquito, length of 
asexual cycle, etc. The morphological differences chiefly 
concern the gametocytes. Those of P. prsecox are more or 
less round, displace the nucleus of the host-cell, and contain 
fine dust-like particles of pigment. The gametocytes of 
P. cathemerium are similar in shape and size, but contain much 
coarser granules of pigment, which tend to have a rod-like 
shape. P. elongatum takes its name from the elongate 
character of the gametocytes, which markedly resemble 
those of Hsemoproteus. Russell (1932) described another 
species. Besides these species, which have been employed 
in experimental work, a large number of other species of 
Proteosoma have been described as distinct, because they 
are found in different species of birds. It is now no longer 
possible to regard the malarial parasites of birds as all belonging 
to the single species P. prsecox ; but the opposite tendency 
towards the multiphcation of species should be avoided 
unless definite morphological distinctions are noted. In 
recent years Ed. and Et. Sergent and Catanei (1931) and 
Giovannola (1934) have attempted a systematic revision of 


species occurring in birds. The latter recognizes nine species 
of avian Plasmodia, and divides them into three groups 
according to their morphology, as follows : — 

(1) Spheroidal gametocytes ; nucleus of the parasitized 
cell displaced by gametocytes and adult schizonts (P. prsecox 
with pointed pigment and P. cathemerium with rod-shaped 
pigment in the gametocytes). 

(2) Elongate gametocytes, spheroidal schizonts ; nucleus 
of the parasitized cell displaced by schizont and sMghtly by 
gametocytes (P. elongatumj. 

(3) Elongate gametocytes ; nucleus of the parasitized 
cell never displaced. This may be divided into two sub- 
groups : 

(a) Adult schizonts small and quadrangular (P. rouxi 

with four merozoites ; P. tenue with 4 to 8 merozoites). 

(b) Adult schizonts circumscribe the nucleus of the para- 

sitized cell (P. circumflexum and P. fallax). 

Coatney and Roudabush (1936) give a list of malarial 
parasites with their hosts, without expressing any opinion 
about the number of valid species. De Mello (1937 a) considers 
the scheme proposed by Giovannola as premature, and refers 
to his description of P. centropi, which combines certain 
characters of all the three groups. 

Buxton (1935) has shown that if females of Culex fatigans 
are infected with Proteosoma, deaths occur earlier than in 
controls, and thinks it due to the invasion of the wall of the 
mid-gut by the ookinete. 

Wolfson (1936) was able to transmit bird malaria by the 
intravenous injection of sporozoites. The salivary glands 
of Culex pvpiens infected with P. prsecox were dissected out 
and the contents placed in a solution of 1 per cent, sodium 
citrate in 0-7 per cent, sodium chloride to which one drop 
of sterile bovine serum per c.c. was added. Canaries were 
injected intravenously with 150 mgm. of the solution, and two 
out of the three birds acquired an infection. 

211. Proteosoma centropi (de Mello). 

^Plasmodium centropi, de Mello, 1937 a, p. 97. 

Gametocytes spheroidal or crescent-shaped ; nucleus of 
the parasitized cell always displaced ; pigment granular 
or rod -shaped. 

Remarks. — Full description of this species has not yet been 
published. According to de Mello (1937 a) the species com- 
bines the characters of all the three groups of Giovannola's 
classification as given above. 

Habitat. — Blood of Centropus sinensis parroti Stresemann : 
Portuguese India. 



212. Proteosoma chloropsidis (de Mello). (Fig. 128.) 

■^Plasmodium prsscox, Scott, 1926, p. 237 ; Wenyon, 1926, p. 1369. 
^Plasmodium chloropsidis, de Mello, 1935 a, pp. 354-5, pi. xliii, fig. 1. 
Plasm.odium, prsecox, Coatney & Roudabush, 1936, p. 342; de Mello, 
1937 a, p. 98. 

Young trophozoites ring-shaped, resembhng those of 
Laverania malariee. Larger trophozoites in the form of big 
rings with brownish pigment granules. Rosettes with (or 
without) pigment, situated generally on the periphery of the 
red cell, with varying number (6 to 11) of merozoites. Male 
gametocytes oval, with large, irregular nucleus. Female 
gametocytes roundish, generally stained deeply blue, with 
round or oval nucleus. Infected red corpuscles not altered. 

Remarks. — De Mello beheves that the form is the same as 
that registered as P. preecox, from the same host, by Scott and 
Wenyon. He, however, regards it as a distinct species, differ- 
ing from true P. prsecox in the irregular number of merozoites 
formed during schizogony, and in the compact structure of 



Fig. 128. 

-Proteosoma chloropsidis (de Mello). A, ring-form; B, full- 
giown trophozoite ; C, rosettes ; D, raale gametocyte ; 
E, F, female gametocytes. (After de Mello.) 

the nucleus of the merozoites, which contrasts with the ring- 
shaped nucleus of the merozoites in P. prsecox. 

Habitat. — Blood of the golden-headed chloropsis, Chloropsis 
aurifrons Temm., from India, in the Zoological Gardens, 
London ; also blood of Chloropsis aurifrons davidsoni Stuart 
Baker : Portuguese India, Mardol (Ponda). 

213. Proteosoma columbse (Carini). 

Plasmodium columbas, Carini, 1912 pp. 396-8. 
Plasmodium sp., Ed. & Et. Sergent, 1904, pp. 132-3. 
•\Hsemoproteus columhae, de Mello & de Sa, 1916, pp. 731-4; de 
Mello, de Sa, de Sousa, Dias, & Noronha, 1917, pp. 8-9. 
Plasmodium columbse, Coatney & Roudabush, 1936, p. 339. 

Schizogony said to take place in the plasma of the blood 
in the lung. Merozoites are seen in red blood- corpuscles as 
small irregular bodies with cytoplasm and nucleus. The 



large schizont reaches nearly a quarter the size of a red cell. 
Stamed by Giemsa's stain the nucleus is bright red and the 
cytoplasm deep blue ; some minute brownish-yellow pigment 
granules can be seen in the latter. It is then said to become 
free in the plasma, lose its pigment, assume a circular form, 
and stain a pale blue, with the chromatin central in position. 
This is called the "Acton body." The chromatin divides, 
forming two, and later more numerous, nuclear masses, some 
of which continue their further division, whilst others change 
into chromidial dust. The "Acton body " grows, and eventually 
aU the nuclei disappear, and in its interior no nucleus, 
but only idio-chromidia, can be seen. This stage, called 
the meroblast, gives origin directly to the merozoites {corps en 
rosace of French authors, roseta of Portuguese authors). The 
roseta breaks up, and the merozoites infect fresh red corpuslces. 

Sexual forms are seen in the peripheral blood. Gamete 
formation, their structure and distinction into maJe and female 
forms, as described by Acton and Knowles for Hsemoproteus 
columhsR. Young gametes can be recognized as male or female 
by their staining reaction, form, situation and arrangement 
of chromatin, and the quantity and size of pigment granules. 
Gametes are halter- shaped, and embrace the nucleus of the 
infected blood- corpuscle. 

Habitat. — Blood of Columha sp. : Portuguese India, Nova 

214. Proteosoma gaUinulse (de MeUo). (Fig. 129.) 

■\ Plasmodium, gallinulse, de Mello, 1935 a, pp. 352-3, pi. xlii, fig. 3 ; 
de Mello, 1937 a, p. 98. 

Young ring-forms very regular. Full-grown trophozoites 
larger in size and oval in form. Large amoeboid trophozoite 
provided with brownish-black pigment set free by the bursting 
of the corpuscle. Gametocytes show a clear sexual dimorphism. 
Male gametocytes are roundish or oval, alveolar ; nucleus 

D E F (5 

Fig. 129. — Proteosoma gallinulse (de Mello). A, ring-form ; B, fuU- 
grown trophozoite ; C, free amoeboid trophozoite ; D, E, 
male gametocytes ; F, G, female gametocytes. (After 
de Mello.) 



consisting of many chromatinic rods irregularly connected 
together ; pigment granules smaller, more or less scattered 
through the cytoplasm. Female gametocytes stain a compact 
and deep blue, have a large nucleus, and the pigment granules 
are larger. Infection very scanty. Infected red cells hyper- 

Habitat. — Blood of the moor-hen, Gallinula chloropus Linn. : 
PoETUGUESB India, lake of CaramboHm. 

215. Proteosoma herodiadis (de Mello). (Fig. 130). 

'fPlasmodiutn herodiadis, de Mello, 1935 a, p. 351, pi. xlii, fig. 1 ; 
de Mello, 1931 a, p. 98. 

Ring form with a granular or rod-like chromatinic dot. 
Older trophozoites pyriform or rounded, with irregularly 
scattered brownish- black pigment. Rosettes mth five to 
ten chromatinic bodies. Male gametocytes somewhat oval, 
with a larger, irregular nucleus. Female gametocytes roundish, 
with a round nucleus. Infection not very heavy. Parasitized 
red cells not altered. 


D E F 

Fig. 130. — Proteosoma herodiadis (de Mello). A, ring-form ; B, C,. 
pyriform or rounded trophozoites ; D, schizont ; E, male 
gametocyte ; F, female gametocyte. (After de Mello.) 

Remarks. — This form is said to be distinguishable from 
Hsemoproteus herodiadis, found in the same host, by the 
nucleus being stained a vivid red colour in the former and 
a pale rose colour in the latter. 

Habitat. — Blood of the heron, Egretta intermedia intermedia 
Wagler : Poetuguese India, lake of Carambolim. 

216. Proteosoma heroni (Basu). 

■f Plasmodium heroni, Basu, 1938, p. 244. 

Remarks — -Morphological description of the species has 
not yet been published. Infection is easily transmitted 
from one paddy-bird to another by blood inoculation, and the 
disease set up is severe. 

Habitat. — Blood of the paddy-bird commonly known as 
" pond heron " : Bengal, Calcutta. 



217. Proteosoma moruony (de Mello & de Sa). (Fig. 131.) 

■\Hazmoprot6us moruony, de Mello & de Sa, 1916, pp. 734-5, pi. Ixiv, 
figs. A, 1-13, B, 1-18; de Mello, de Sa, de Sousa, Dias, & 
Noronha, 1917, p. 9. 

Hasmoproteus moruoni, Wenyon, 1926, p. 1370. 

Hce')noproteus moruony, de Mello, 1937 a, p. 99. 

Schizogony said to take place in the plasma. Merozoite 
enters a red blood- corpuscle and assumes a circular form, 
with a small chromatinic mass in the centre. Pigment granules 
are present. Large schizonts not seen in the body of ery- 
throcytes, and the largest form in a red cell does not reach 
a quarter of the size of the infected cell. The schizont becomes 
free in the plasma, as the "Acton body." It has no visible 
pigment, and is surrounded by a very dehcate hyahne capsule. 
The chromatin undergoes binary division, forming eight 
chromatin dots. Later, all the chromatin changes into idio- 
chromidia and the capsule eventually disappears, the mero- 
zoites constituting the roseta. 

Gamete formation, their structure, and differentiation into 
male and female similar to that in Hsemoproteus columbse. 

Fig. 131. — Proteosom,a m.oruony (de Mello & de Sa). A, young tropho- 
zoite; jB, full-grown trophozoite ; C, "Acton body," show- 
ing nuclear division ; D, schizont ; E, rosetta containing 
merozoites. (After de Mello and de Sa.) 

The nucleus in an adult gamete is situated in the convexity 
of the halter and nearer to one pole than the other. The 
chromatin of the macrogamete is described as undergoing 
division for regressive schizogony. 

Remarks. — De Mello and de Sa, though describing this 
parasite as a Hsemoproteus species, remark that all the stages 
of its development are seen in the peripheral blood, and no 
special forms are to be found in the internal organs. 

Habitat. — Blood of the magpie robin, Gopsychus saularis 
(Linn.) : from India, in the Zoological Gardens, London ; 
also from the same host : Portuguese India, Calangate 

218. Proteosoma prsecox (Grassi & Feletti). (Fig. 132.) 

Hsemam,aiba prmcox, Grassi & Feletti, 1890, p. 463. 
H3em,am,ceba relicta, Grassi & Feletti, 1890, p. 465. 
H3em,oproteus danilewskii (part), Kruse, 1890, p. 371. 
Hsemoproteus, vars. b & c, Celli & San Felice, 1891, pp. 541-8, 
pis. vii, viii. 



Proteosoma grassii, Labbe, 1894, p. 157, pi. ix, figs. 1-31. 
'\Prot60soma sp., Ross, 1898, pp. 401-8, 448-50 ; 1899 a, pp. 1-3 ; 

1899 b, pp. 136-44 ; Daniels, 1899, pp. 443-54. 
Heemproteus danilewskyi, Labbe, 1899, p. 80, fig. 146. 
Proteosoma sp., Schaudlnn, 1900, pp. 159-81 ; Grassi, 1900, 

pp. 115-24 ; Ruge, 1901, pp. 187-91, 2 figs. ; Hartmann, 1907, 

pp. 148, 149, 152. 
Plasmodium praecox, Minchin, 1912, p. 358. 
Plasmodium, relictum, Minchin, 1912, p. 358. 

Proteosoma prsecox, Doflein, 1916, pp. 897-901, figs. 896-900, 903. 
^Proteosoma prsecox, de Mello, de Sa, de Sousa, Dias, & Noronha, 

1917, p. 15. 
■\Plasmodium danilewskyi, Castellani & Chalmers, 1919, p. 513. 
Proteosoma praecox, Calkins, 1926, p. 444. 
^Plasmodium prsecox, Scott, 1926, p. 237 ; Wenyon, 1926, pp. 1365, 

Plasmodium prsecox, Wenyon, 1926, pp. 977-8, pi. vi, figs. 1-10 ; 

Hegner, 1926, pp. 479-80 ; Hartmann, 1927, pp. 1-7, pis. i, ii ; 

Knowles, 1928, pp. 442-3, fig. 102, 1-i ; Hegner, 1929, pp. 59- 

Proteosoma prsecox, Reichenow, 1929, pp. 983-4, figs. 952-4, 1000. 
Plasmodium prsecox, Manwell, 1930, pp. 382-3 ; Kudo, 1931, 

p. 287, fig. 122 a-d. 
Plasrnodium relictum, Ed. & Et. Sergent & Catanei, 1930, pp. 223- 

38 ; 1931 a, pp. 399-437 ; 1931 h , pp. 327-35. 
Plasmodium prsecox, Giovannola, 1934, pp. 372—8, pi. i, figs. 1—4 ; 

Coatney & Roudabush, 1936, p. 340 ; de Mello, 1937 a, p. 95. 

Schizogony takes place in the red blood-corpuscles, generally 
in the viscera. Young trophozoites resemble the small rings 


Fig. 132. — Proteosoma prsecox {(jTassi &, Yeletti). (x2000.) J., young 
ring-shaped trophozoite ; B, G, larger trophozoites ; 
D, schizont splitting into a number of merozoites ; E, female 
gametocyte ; F, male gametocyte. (After Wenyon.) 

of Laverania malarise. As growth proceeds, dark pigment 
granules appear, and the host- cell becomes distorted, its 
nucleus becoming displaced more or less to one side or to 
one end of the cell (unlike Hsemoproteus, which develops in 
the red blood- corpuscle without deforming it or displacing 
the nucleus to any extent). Schizont varies in size and 
period of growth. Small schizonts measure 4-5 /x, and 
produce only about six merozoites. Largest schizonts occupy 
about half the corpuscle and produce 16 to 24 merozoites. 


the average number of merozoites formed being 15 ; in such 
cases the infected corpuscles are altered in shape, pale in 
colour, and have the nucleus displaced to one side of the 
corpuscle. Sometimes two or more schizonts occur in a single 
cell, and a crop of as many as 36 merozoites may result. 
In the fully developed schizont the pigment is seen as a single 
dark mass. Gametocytes are sHghtly elongate ovoid bodies 
about the size of the largest schizonts, and closely resemble 
those of Laverania. The female gametocyte stains more 
deeply and has a more compact nucleus than the male. 
Infection is transmitted from bird to bird by Cidex mosquitoes. 
Sporogony is similar to that of the human malarial parasites 
in Anopheles, the time occupied by the cycle varying with 
temperature. Number of oocj^sts varies with the number of 
gametocytes in the blood, in some cases the whole wall of the 
stomach of the mosquito being studded with the oocysts. 

The parasite usually occurs in the blood of small birds like 
the sparrow and lark, but can also be found in the blood of 
larger birds such as the crow, owl, pigeon, partridge, duck, fowl, 

Remarks. — ^The species is of special interest, as it was while 
investigating it that Ross (May, 1898) first discovered the 
stages of fertihzation and oocyst formation in the stomach 
of Culex sp. (probably C. fatigans Wied.). In June and July 
of the same year Ross discovered the liberation of sporo- 
zoites into the coelomic cavity of the mosquito and the invasion 
of the salivary glands, and succeeded in infecting healthy 
birds from infected Culex mosquitoes. He thus discovered the 
whole of the transmission cycle in P. prsecox. He had previously 
(1897) discovered oocysts of human malarial parasites in 
Anopheles mosquitoes, and predicted that the transmission 
cycle of human malarial parasites would be found to occur in 
Anopheles mosquitoes, and that it would be identical with that 
of P. precox in Culex, thus paving the way for the discovery 
of the complete hfe- cycle of the human malarial parasites 
by the Italian workers. 

Daniels (1899), Avorking at Calcutta, confirmed Ross's 
observations. Koch, working in Italy (1899 a), studied the 
development in Culex nemorosus {=Aedes nemorosus), and 
Grassi (1899) in C. pipiens. Liihe (1900) clearly brought out 
the relation of the hfe- cycle of malarial parasites to that of 
CocciDiA, and introduced a terminology for various stages 
that is in vogue till to-day. Ed. and Et. Sergent (1907) and 
Neumann (1908) demonstrated that the complete cycle 
could take place in a small proportion of Aedes segypti (Linn.) 
{=^Stegomyia fasciata) that were fed on infected blood. Ed. 
and Et. Sergent showed later (1918) that the development 
could take place in certain other species of Aedes and Culex 


also, and they (1919) carried out experiments on the develop- 
ment of P. prsecox under varying conditions of temperature. 

Celli and San Felice (1891) were the first to show that the 
infection could be transmitted from lark to lark by inoculation 
of blood from an infected bird to a healthy one. Grassi and 
Peletti (1891) and Laveran (1891) repeated the experiment 
in larks, and Ziemann (1898) in another type of bird. Ross 
(1898 6) showed that mosquitoes fed on sparrows were able 
to infect crows and larks. Koch (1899 a, 1899 6) showed 
that parasites naturally occurring in one type of birds can be 
inoculated into other types of birds. Ruge (1901) and 
Wasielewski (1908) were similarly able to infect various types 
of birds, thus showing that the same species of Proteosoma 
is able to infect a number of different genera and species. 
According to HuiBP (1927) infections had been noted in at least 
seventy-nine species of birds belonging to at least fifteen 
famihes. Originally the malarial parasites of birds were all 
considered as belonging to a single species, but they have now 
been spht up into a number of species by Hartmann (1927), 
Huff (1930), Manwell (1930) and others, and parasites from 
a large number of different birds have also been described as 
distinct species. Mayne (1928) has recorded both natural and 
experimental infections in Anopheles subpictus Grassi {=^rossii). 
According to Giovannola (1934) no less than four species are 
known to occur in Passer domesticus (Linn.), viz., P. precox 
Grassi & Feletti, P. cathemerium Hartmann, P. elongatum Huff, 
and P. rouxi Ed. & Et. Sergent & Catanei. According to him 
P. prsRCox is distinguishable by spheroidal gametocytes con- 
taining pointed pigment granules, and spheroidal schizonts 
producing 14 to 32 merozoites. 

In recent years a good deal of experimental work has been 
carried out on infection and resistance in bird malaria (Boyd, 
1925 ; Taliaferro, 1925 ; Hegner, 1926), on relapses in bird 
malaria (Manwell, 1929), and on quinine and plasmochin 
therapy (Boyd, 1926 ; Hegner, Shaw, and Manwell, 1928 ; 
Manwell, 1930), resulting in the discovery of facts which 
are of considerable practical importance in the warfare against 
human disease. 

" Black spores " of Ross. — Certain bodies described as 
" black spores " were originally noted by Ross (1899, 1905, 
1923) and Daniels (1900) within oocysts in the stomach of 
infected mosquitoes, as well as free in the tissues, and even in 
uninfected mosquitoes. According to Stephens and Christo- 
phers (1908) they are also found in the thoracic muscles and 
salivary glands. Castellani and Chalmers (1919) regard these 
so-called " black spores " as protozoal parasites of the genus 
JVosema, which have invaded the oocyst. 


Bruce Majme (1929), after reviewing the previous literature 
on the subject, and after recording his observations to the 
effect that these " black spores " have been found during 
experiments with avian malaria in uninfected mosquitoes, as 
well as in mosquitoes harbouring the parasites of malaria, 
and even in freshly emerged, unfed, laboratory-bred females and 
males of Anopheles, Gulex, and Musca, comes to the conclusion 
that most, if not all, " black spores " appear to be merely 
chitinogenous thickenings of the tracheal tubes. Eaiowles and 
Basu (1933) conclude that no less than three, if not four, 
different structures have been described as " black spores " 
or " chitin corpuscles " by different workers. These are : 
(a) True malaria oocysts which have undergone degeneration, 
and in which a very heavy deposition of pigment has occurred ; 
these alone they beheve to be the true " black spores " 
of Ross ; (b) a hyper-chitinization of segment-like portions of 
the finer ramifications of the tracheal system ; (c) fungus 
infections of the tracheal system ; (d) infections of the mos- 
quito tissues concerned with miscrosporidian parasites. 
Their observations, based upon material studied in sections, 
led them to conclude that (b), (c), and (d) have nothing to do 
with true " black spores," which had lietter be described as 
" degenerated and hyper-pigmented oocysts " or ruptured 
contents of the same. 

Habitat. — Blood of sparrows, larks, pigeons, crows : Bengal, 
Calcutta ; some birds : Ceylon ; oocysts in mosquitoes, Gulex 
fatigans Wied., fed on these birds ; also in Anopheles subpictus 
Grassi (=ro55n) : also blood of white-throated munia, Uroloncha 
malabarica Linn., and common Indian starKng, Sturnus 
vulgaris poltaratshyi Finsch, from India, and Tragopan satyra 
(Linn.), from the Himalayas, in the Zoological Gardens, 

219. Proteosoma sp. 

^Plasmodium (?) sp., Scott, 1926, p. 237 ; Wenyon, 1926, p. 136 6. 
Plasmx)dium sp., Coatney & Roudabush, 1936, p. 341 ; de Mello, 
1937 a, p. 95. 

Habitat. — Blood of the sarus crane, Antigone antigone (Linn,), 
from India, in the Zoological Gardens, London. 

220. Proteosoma sp. 

^Plamnodium, sp., Donovan (first recorded in Wenyon, 1926, 
p. 1371). 
Plasm,odium sp., de Mello, 1937 a, p. 95. 

Habitat. — Blood of the Indian house crow, Gorvus svle.nde.ns 
Vieillot : India (locaUty not cited). 

SPOK. s 


221. Proteosoma sp. 

^Plasmodium sp., Plimmer, 1914, p. 190. 

Plasmodium sp., Wenyon, 1926, p. 1371 ; Coatney & Roudabiish, 
1936, p. 342. 

Habitat. — Blood of the barbet, Cyanops flavifrons (Cuvier), 
from Ceylon, in the Zoological Gardens, London. 

222. Proteosoma sp. 

^Plasm,odium, sp., Plimmer, 1913, p. 148. 
Plasm,odium, sp., Wenyon, 1926, p. 1372 ; Coatney & Roudabush, 
1936, p. 342 ; de Mello, 1937 a, p. 95. 

Habitat. — Blood of the grey- headed bunting, Emberiza fucata 
Pallas, from India, in the Zoological Gardens, London. 

223. Proteosoma sp. 

^Plasmodium sp., Plimmer, 1912, p. 415 ; Scott, 1925, p. 237 ; 
Wenyon, 1926, p. 1372. 
Plasmodium sp., Coatney & Roudabush, 1936, p. 342 ; de Mello, 
1937 a, p. 95. 

Habitat. — Blood of the wattled starHng, Gracula indica 
(Cuvier) [=Eulabes religiosa Auct.],from South India, in the 
Zoological Gardens, London. 

224. Proteosoma sp. 

■\Plasm,odium, sp., Plimmer, 1912, p. 415. 

Plasmodium, s^., CoatuBj & Roudabush, 1936, p. 342; de Mello, 
1937 a, p. 95. 

Habitat. — Blood of the jay-thrush or babbler, Garrulax 
leucolophus (Hardwicke), from North India, in the Zoological 
Gardens, London. 

225. Proteosoma sp. 

■\Plasmodium sp., Scott, 1926, p. 237, Wenyon, 1926, p. 1377. 
Plasm,odiutn sp., de Mello, 1937 a, p. 95. 

Remarks. — Laveran and MaruUaz (1914) have described 
Proteosoma [Hsemamceba) tenue and Proteosoma [Hsemamceba) 
liothiris from the same host from Japan. 

Habitat. — Blood of the babbler, LeiotJirix lutea (Scop.), from 
India, in the Zoological Gardens, London. 

226. Proteosoma (?) sp. 

■\Plasmodium (?) sp., Scott, 1926, p. 237 ; Wenyon, 1926, p. 1377. 
Plasm,odium, sp., Coatney & Roudabush, 1936, p. 343. 

Habitat. — Blood of the Ceylon loriquet, Coryllis beryllinus 
(Forster) [=Loriculus indicus Gmel.], from Ceylon, in the 
Zoological Gardens, London. 


227. Proteosoma sp. 

■fPlasmodium sp., Plimmer, 1913, p. 148. 
Plasmodium s-p., Wenyon, 1926, p. 1377 ; Coatney & Roudabush, 
1936, p. 343 ; de Mello, 1937 a, p. 95. 

Habitat. — Blood of the bunting, Melophus mdanicterus 
(Gmel.), from India, in the Zoological Gardens, London. 

228. Proteosoma ap. 

^Plasmodium sp., Plimmer, 1915, p. 130. 
Plasmodium sp., Wenyon, 1926, p. 1377 ; de Mello, 1937 a, p. 96. 

Habitat. — Blood of the thrush, Turdus boulboul (Latham), 
from India, in the Zoological Gardens, London. 

229. Proteosoma sp. 

■fPlasmodiutn sp., Plimmer, 1912, p. 415. 
Plasmodium sp., Wenyon, 1926, p. 1383 ; de Mello, 1937 a, p. 96. 

Habitat. — Blood of the bulbul, Elathea jocosa (Linn.) 
[:=Otoco'mpa eimeria], from India, in the Zoological Gardens, 

230. Proteosoma sp. 

■fPlasmodium sp., Plimmer, 1913, p. 148. 
Plasmodium sp., Wenyon, 1926, p. 1382. 

Habitat. — Blood of the chat, Saxicola caprafa (Linn.), from 
India, in the Zoological Gardens, London. 

Genus LAVERANIA Grassi & Feletti, 1890. 

A third type of malarial parasite, Golgi, 1886 a, p. 109 ; 1886 6, 

p. 419; 1889, p. 173. 
Malignant tertian parasite, Marchiafava & Colli, 1889. 
Laverania, Grassi & Feletti, 1890, p. 4 ; 1892, p. 10. 
HcBmatomonas, Hehir, 1893, pp. 207-12, 241-8, 273-80. 
Hcem,atozoon, Welch, 1897. 
Plasmodium (part), Labbe, 1899, pp. 80-2; Liihe, 1900, p. 392; 

Schaudinn, 1902. 
Laverania, Neveu-Lemaire, 1900, p. 9; Minchin, 1903, pp. 243, 

Hsemam,oeba (part), Laveran, 1907, pp. 110-60. 
Plasmodium, (part), Minchin, 1912, p. 358. 
Laverania, Doflein, 1916, pp. 902-12 ; Castellani & Chalmers, 1919, 

pp. 517-18. 
Plasmodium (part), Muhlens, 1921, pp. 1502-11. 
Laverania, Thomson & Woodcock, 1922, pp. 1530-4. 
Plasmodium (part), Hegner & Taliaferro, 1924, pp. 327-9 ; Wenyon, 

1926, pp. 934-41 ; Knowles, 1928, pp. 392-400. 
Laverania, Stiles, 1928, p. 881 ; Reichenow, 1929, pp. 987-98. 
Plasmodium (part), Kudo, 1931, p. 286 ; Calkins, 1933, pp. 406, 

407, 566. 
Laverania, Reichenow, 1935, p. 377. 

Plasmodium (part), Brmnpt, 1936, pp. 418-24 , Coatney & Rouda- 
bush, 1936, pp. 338-53. 

Trophozoites very small rings. Schizonts also remain small 



and do not completely fill the red-blood corpuscles. Hsemo- 
zoin pigment granular. Gametocytes elongated, wormlike- 
and somewhat sickle-shaped, so-called " crescents." Parasites 
of man and anthropoid apes. 

Remarks. — Laverania malaria, the parasite of mahgnant 
tertian or tropical epidemic malaria in man, and Laverania 
reichenowi, the parasite of the gorilla and chimpanzee, are 
included in this genus. The species of 'this genus show 
considerable resemblance with Proteosoma of birds. 

231. Laverania malarise Grassi & Feletti*. (Figs. 122 (PL I), 
133, & 134). 

Malarial parasites, Laveran, 1880 a, p. 158; 1881, pp. 627-30; 

1882, p. 737. 
Oscillaria malarise (part), Laveran, 1883, p. 113. 
A third type of malarial parasite, Golgi, 1886 a, p. 109 ; 1886 b, 

p. 419; 1889, p. 173. 
Hsemamceba malarise, Laveran, 1890, p. 374. 
Hsemamoeba malarise prsecox-\-H. malarias immaculatum, Grassi & 

Feletti, 1890, p. 10. 
Hsemamceba prsecox, Grassi & Feletti, 1890, p. 6 ; 1892, p. 10. 
Laverania malarise, Grassi & Feletti, 1890, p. 4 ; 1892, p. 10. 
Hsemamoeba febris quotidinse, Marchiafava & Bignami, 1891. 
Hsemamceba immaculata, Grassi, 1891, p. 14; Grassi & Feletti, 

1892, p. 10. 
■[Hsematomonas malarias, Hehir, 1893, pp. 207-12, 241-8, 273-80, 
pis. i-viii. 
Hsematozoon falciparum, Welch, 1897, pp. 36, 47. 
Plasmodium malarise prsecox, Labbe, 1899, p. 82. 
Plasmodiujn malarise iinmaculatum, Labbe, 1899, p. 82. 
^Hsemomenas prsecox, Ross, 1899, pp. 322-4, 439 ; 1900, pp. 522-7. 
Plasmodium prsecox, Liihe, 1900, p. 460. 
Laverania prsecox, Neveu-Lemaire, 1900, p. 9, pi. i, fig. 37. 
Laverania malarise, Minchin, 1903, pp. 243-54, 267, 332, 351, 

fig. 68. 
Hsemamoeba malarise, Laveran, 1907, pp. 110-60 ; fig. ii, pi. i. 
Plasmodium prsecox, Doflein, 1909, pp. 662-70, figs. 596-609. 
Laverania malarise, Doflein, 1909, pp. 662-70, figs. 596-609. 
Plasmodium falciparum, Bass & Johns, 1912, pp. 567-79 ; Minchin, 

1912, pp. 358-60, fig. 156. 
Malignant tertian malarial parasites, Thomson & Thomson, 1913, 

pp. 77-87, pi. X. 
^Plasmodium falciparum. Row, 1917, p. 392, pi. xxiii. 
■\Lav6rania prsecox, Row, 1917, p. 392, pi. xxiv. 
Laverania malarise, Castellani & Chalmers, 1919, pp. 517-18, figs. 

174, 175, pi. i, figs. lc-4c. 
Plasmodium immaculatum, Miihlens, 1921, pp. 1502-11, pi. xxxi, 

figs. 20-50 ; pi. xxxii, figs. 26-40 ; pi. xxxiii, figs. 2, 5-21 ; 

text-figs. 4-7. 

* The retention of the gemis Laverania for the parasite of malignant 
tertian Malaria is in accordance with Opinion 104 of the Intern. Conam. 
on Zool. Nomenclature. Many malariologists, however, consider that 
the separation of this parasite from those of benign tertian and quartan 
Malaria is not required on zoological grounds, and include all three 
species in the geuus Plasmodium ; if this latter classification be followed, 
the name of the malignant tertian parasite will be Plasmodium falci- 
parum (Welch). — Editor. 


Laverania malarise, Thomson & Woodcock, 1922, pp. 1530-6, 

pis. Ixv, Ixvi ; text-figs. 545, 546. 
Plasinodium falciparum, Hegner & Taliaferro, 1924, pp. 327—9, 

pi. ii, figs. 9-16. 
Laverania falciparum. Stiles & Hassall, 1925, p. 42. 
Plasmodium falciparum, Craig, 1926, pp. 433-52, figs. 74-8 ; Wen- 
yon, 1926, pp. 934-41, pis. vii-ix, pi. xiii, figs. 16-40 ; figs. 391, 
401-3; Hehir, 1927, pp. 166-73, pi. xii, figs. 76-117; pi. xiii, 
figs. 118-29 ; pi. xiv, figs. 1-8 ; figs. 59-60. 
'^Plasmodium falciparum, Knowles, 1927, pp. 14-20, pi. iii, figs. 1-24 ; 
1928, pp. 392-400. pi. xiii. 
Laverania malarix, Reichenow, 1929, pp. 987-97, figs. 958-73, 
975 a-e. 
'^Plasmodium falciparum. Row, 1929, pp. 1120-5, pis. Ixxxviii, 
Ixxxix ; 1930, pp. 221-6, pis. xxiii, xxiv. 
Plasmodium' falciparum, Aragao, 1930, pp. 41-56, 1 pl. ; Kudo, 
1931, p. 286, fig. 121, h-n ; Calkins, 1933, p. 407, pl. i, figs. 13-17 ; 
pl. ii, figs. A-L ; Chaoulitch, 1936, pp. 716-22 ; Brumpt, 1936, 
pp. 418-24, figs. 185-7 ; Coatney & Roudabush, 1936, p. 339. 
Plasmodium immaculatum, Giovannola, 1935, pp. 90-1. 
^Plastnodium tenue, Stephens, 1914, pp. 169—72. 

The so-called Plasmodium tenue, Balfour & Wenyon, 1914, p. 353. 
■fPlasmodium tenue, Sinton, 1922, pp. 215-35, pis. ii & iii ; Knowles, 
1923 a, p. 276 
Plasmodium tenue, Wenyon, 1926, pp. 951-2, fig. 405 ; Craig, 1926, 

pp. 464-70. fig. 81. 
'\Plasmodium tenue, Kjiowles, 1928, p. 404, fig. 93. 
Plasmodium tenue, Reichenow, 1929, p. 988, fig. 961. 

The Cycle in Man. — Young trophozoites are very small, 
often seen as a tiny bead of chromatin, "with a wisp of blue- 
staining cytoplasm adhering to the margin of the red corpuscle 
(the marginal or accole form). Multiple infection of red 
blood- corpuscles is very common, many of them containing 
two, three, or sometimes a larger number. The smallest 
" rings " are very narrow, appearing as a thin blue hne 
surrounding a vacuole, with a red-staining granule of chromatin 
or nucleus protruding at one side, occupying no more than 
one-sixth of the diameter of the red blood- corpuscle. Chroma- 
tin often rod-like, and divides very early into two granules 
(nuclei). Fully grown trophozoites may be as large as half 
the size of the red blood-corpuscle, and the cells containing 
these big rings show Maurer's dots. Occasionally more 
irregular or amoeboid forms occur. Infection is usually 
heavy, as many as 25 per cent, of the corpuscles being found 
infected. Infected red corpuscles do not become enlarged 
(as they do in infections with Plasmodium vivax), but on their 
surface appear red- staining markings usually known as 
Maurer's dots (the markings were actually first described 
by Stephens and Christophers, who regarded them as clefts 
in the red corpuscle in which the red of the stain is deposited) . 
Maurer's dots are larger and fewer in number than Schiijffner's 
dots (which are found in P. vivax), and are best seen in deeply 
stained films : they are stellate or crack-hke rather than dot- 
like, and stain brick-red rather than the bright pink of 

Fig. 133. — Life-cycle of Laverania malarial Grassi & Feletti in man 
and the mosquito. ( X c. 1000.) A-G, schizogony in 
red blood-corpuscles ; H, immature gametocyte ; I^, 1^, 
male and female gametocytes in blood of man ; J^-L^, 
development of male gametocyte and production of 
microgametes ; J^-L^, development of female gameto- 
cyte and extrusion of polar body in the stomach of mos- 
quito ; M, fertilization ; N, transformation of zygo te into 
ookinete which penetrates through the intestinal epithelium ; 
O, P, development of oocyst between the epithelium and 
the elastic membrane ; Q-S, growth of cytoplasm accom- 
panied by multiplication of nuclei and vacuolation of 
cytoplasm ; T, vacuoles run together to reduce cytoplasm 


Schiiffner's dots. Sooner or later hsemozoin pigment appears 
in the growing trophozoite, the pigment being dark brown or 
black, and is collected into a distinct round mass, and this 
early collection of the pigment into a round compact mass 
in the preschizogonic stages is a diagnostic feature of the 
asexual forms of this species. 

All the infected corpuscles soon disappear from the peri- 
pheral circulation, and schizogony takes place almost exclusively 
in the internal organs. Sometimes this takes place before 
pigment has formed, at others not till some growth has taken 
place and pigment granules have developed in the cytoplasm. 
It was probably this that led the earher observers to describe 
two separate species of malignant tertian parasities (P. malarise 
'prsecox and P. malarise, immaculatum) . The pigment granules 
run together to form a dark granular mass at one side of the 
parasite. The fully developed schizont occupies one-half to 
two-thirds of the diameter of the red blood-corpuscle, and gives 
rise to 8 to 24, usually 10 to 20, very small merozoites. Sometimes 
a larger number, up to 32, are set free, but this may be due to 
two schizonts being contained in a single corpuscle. The 
merozoites are arranged in a grape-like cluster (" rosette ") 
round a central residual mass, with a considerable amount of 

The attack of malaria synchronizes with the setting free of 
the merozoites, but the three stages of the attack are not so 
sharply defined as in the case of P. vivax infections. The fever 
may occur every forty-eight hours (tertian) or every twenty-four 
hours (quotidian) . In the latter case it is probably due to two 
infections having originally taken place, schizogony due to 
each taking place on alternate days, causing a daily attack of 
fever. In certain cases of heavy infections schizonts are seen 
in the peripheral blood along with the trophozoites ; and this 
is usually an indication that the infection is a very serious 
one and may prove fatal. Schizogony may be seen in a certain 
number of ordinary infections in thick blood-films ; but it 
can, however, be studied best in cultures or by carrying out 
spleen punctures during the acute phases of the attack. 
After schizogony, which, as remarked above, usually takes 
place in the internal organs, ring-forms appear again in the 
peripheral blood. 

to a coarse network, with sporozoites commencing to 
form as finger-like buds ; U, sporozoites fully formed and 
attached to several masses of cytoplasm into which the 
network has broken, two residual masses containing 
nuclear matter also present ; V, detached sporozoites 
irregularly distributed in the oocyst, which ruptm-es, 
liberating them into the ccelomic cavity ; W, sporozoites 
entering the salivary glands ; X, sporozoites injected 
into man by the mosquito : they invade the red blood- 
corpuscles and start the schizogony cycle. (After Wenyon.) 


Merozoites grow into gametocytes almost exclusively in the 
blood- stream in the internal organs . Only matm-e gametocytes 
are generally seen in the peripheral blood, though occasionally 
yoiuig forms may also be found. Young gametocytes 
are shghtly elongate, with one side concave and the other 
convex. Fully developed gametocytes are crescentic or 
sausage-shaped, and are generally referred to as " crescents." 
They are about one and a half times the length and about 
half the breadth of a red corpuscle. They usually have rounded 
ends, but sometimes the ends are pointed, and the "crescents" 
are then sickle-shaped. In deeply stained films the margin 
of the blood- corpuscle is seen to be closely apphed to the 
convex side of the crescent and stretched across the concave 
side to show a bulge. Sometimes the crescent appears to 
be completely surrounded by a somewhat irregular red-staining 
border, which J. D. Thomson (1917) interpreted as a definite 
capsule. Two types of crescents are found. The male, or 
microgametocyte, is shorter and broader than the female, 
and has more rounded ends ; it has hyahne cytoplasm, which 
stains a faint blue or a faint dirty pink colour with Romano wsky 
stains ; the nucleus is larger and paler when stained, and 
the pigment is irregularly distributed throughout the middle 
two-thirds of the crescent. The female gametocyte is thinner 
and longer and has more pointed ends ; its cytoplasm is denser 
and stains more deeply blue ; the nucleus is compact and stains 
more intensely, and the pigment is aggregated in the centre 
of the crescent round the nucleus. In well-stained, wet- 
fixed fihns the nucleus of the female gametocyte is seen to 
have a large karyosome, while that of the male has fine chroma- 
tin granules distributed through it. The number of crescents 
varies considerably ; sometimes a single crescent will be found 
after a prolonged search, at other times they are so numerous 
that one or more will be found in nearly every field. Relapses 
are less common than in the benign tertian fever, and the 
infection as a rule dies out more quickly. 

The Cycle in the Mosquito. — Ab in the other species of human 
malarial parasites, the gametocytes undergo their further 
development in the stomach of Anophehne mosquitoes. 
There the crescents retract to a spherical form, thus causing 
the rupture of the remains of the red blood-corpuscles, and 
escaping out of them. Subsequent development and fertihza- 
tion takes place as in the other species. The male gametocyte 
gives rise to a number of microgametes by ex-flagellation, and 
the female gametocyte forms a single macrogamete. Macro- 
and microgametes unite in pairs, and zygotes, called ookinetes, 
are formed, which, after penetrating the stomach-wall, 
become lodged between the epithehum and the elastic layer of 
the wall of the stomach. There they grow ; the nuclei undergo 



rapid and repeated divisions, and finally produce an enormous 
number of minute sporozoites. These sporozoites are set free 
through the rupture of the cyst-wall into the body-cavity, 
find their way into the salivary glands, and are inoculated 
into a new victim when the mosquito bites one. The ookinetes 
and the developing oocysts of this species are distinguished 

m t 

Fig. 134. — Oocysts of Laverania malarise Grassi & Feletti projecting 
from the wall of the stomach of an infected Anopheles; 
c, crop ; m.t., Malpighian tubules ; o, oocysts of Laverania ; 
T, ventral reservoir ; s, stomach, the enlarged portion of the 
mid-gut ;, salivary glands. (From Reichenow, after 
Ross and Grassi.) 

from those of P. vivax by the dark colour of the pigment m 
the former and the light brown or yellow pigment in the 
latter. Also a much larger number of oocysts occurs, and they 


may be so numerous as to be actually in contact with one 
another on the stomach-wall. 

Remarks. — There is a vast literature on this, as well as the 
other species causing malaria in man, and it is difficult to 
refer to even the more important works here. Sinton (1929) 
has pubhshed a very valuable and complete bibliography 
on malaria in India, indexing about 2200 papers and reports 
dealing with the causation, prevention, and treatment of the 
disease. Covell (1927, 1931) has similarly collected all the 
available data relating to the transmission of malaria by 
different species of Anophehne mosquitoes. For information 
on the subjects of pathology of malaria infections, the occur- 
rence and mechanism of relapses, the factors which influence 
the development and survival of malarial parasites in 
mosquitoes, the possibility of animal reservoirs and the 
susceptibility of man and animals to inoculation, reference 
may be made to text-books and other special works. 

L. malarise is usually present in tropical countries, and is 
responsible for severe outbreaks of epidemic malaria. The 
fever is referred to as malignant, as it does not yield readily 
to treatment. Two types are kno\^Ti, a quotidian, which 
recurs every day, and a tertian, which recurs every other day. 
Whether or not these are due to distinct species or the 
quotidian is due to double infection is still uncertain. 

Knowles (1919) noticed crescents, without asexual forms, 
in blood -films taken from a convalescent malaria patient. 
It is quite common to come across crescents and other forms 
in blood- films from persons Kving in an endemic area who are 
quite free from all symptoms of malaria at the time. Knowles, 
Chopra, Gupta, and Das-Gupta (1923) record the examination 
of twelve villagers from the Kuki Hills in Assam admitted to 
hospital in Calcutta suffering from framboesia. All except 
one were afebrile, and yet blood examination showed that 
eleven out of twelve patients harboured malarial parasites, 
all three species being found among different patients, and both 
trophozoites and gametocytes were encountered. Sinton 
(1926) has made a thorough study of the problems relating 
to gametocyte formation. He finds that (a) there is a marked 
seasonal variation in crescent production in patients ; (6) differ- 
ent localities show a different proportion of crescent- carriers ; 
(c) there is a distinct correlation between the numerical pre- 
valence of asexual forms in the peripheral blood and the number 
of crescents which appear about ten days later ; {d) the 
development of crescents appears to be associated with a 
lowered immunity, possibly due to a change in the reaction 
of the pulp in the bone-marrow and spleen ; (e) no marked 
correlation can be traced betAveen the degree of splenic enlarge- 
ment and the occurrence of crescents in the peripheral blood ; 
(/) duration of life of a crescent may be as long as forty to 


fifty days in the peripheral blood, but the great majority of 
them disappear within three weeks after the asexual cycle 
has been destroyed ; and (g) the reduction in the number 
of crescent carriers by efficient treatment is an important 
factor in all anti-malarial campaigns. 

Row (1929), discussing the subject of evolution of the 
crescents of malarial parasites grown anaerobically in simple 
cultures, recorded definite variations during different attacks of 
the same infection. Thus, when a man fresh from Europe, and 
otherwise healthy, is infected, one may find in his first paroxysm 
a very severe cHnical reaction, and yet the number of parasites 
in his peripheral blood is very small. These, however, yield 
a larger number of merozoites in culture, whereas during the 
second and third paroxysms, though the number of parasites 
in his peripheral blood may be very large, the number of 
merozoites yielded in culture is much smaller, and this diminu- 
tion progressively continues for each paroxysm until a time 
is reached when one merozoite entering a red blood- corpuscle 
produces no more than one individual. This is the point 
when the crescent and gamete production is initiated, and, once 
started, it continues, so as ultimately to flood the blood with 
these resistant forms of the parasite, which persist for several 
weeks during the apparent well-being of the patient. The 
factors contributing to this change in the direction of parasitic 
development have been observed to depend {a) on the vigour 
of the phagocytes, and (b) on the biochemical action of the 
plasma or serum in which the free merozoites find themselves 
either after the rupture of the fully developed schizont or 
after their escape from the phagocytes or on (a) and (b) com- 
bined. Row described the phagocytosis in leucocytes and 
plasmolysis by blood serum of such of the merozoites as are 
Hberated by degenerating leucocytes. In vivo these liberated 
merozoites probably initiate a fresh paroxysm. The merozoites 
during each succeeding paroxysm become more resistant 
owing to their passage through the leucocytes, but lose their 
capacity to multiply, until the point is reached when one 
merozoite yields but one individual for each corpuscle attacked, 
and the crescent formation is inaugurated and continued. 
This is an ideal state of equilibrium between the host and the 
parasite. Row also described the actual development of cres- 
cents in cultures, and advanced evidence for the behef that 
antibodies are formed in a malarial infection. Row (1930) 
further showed that an accelerated effort toward crescent 
formation was brought about by using for culture a mixture 
of Laverania malarim and Plasmodium vivax, and put forward 
the suggestion that in nature also such mixed infections may 
be one of the factors responsible for the presence of the 
crescents in carriers in endemic areas. 


Aragao (1930) has studied the development of the gameto- 
cytes of Laverania malar im, and finds that there are two 
types of merozoites, corresponding to the male and the female 
gametocytes. Merozoites destined to form male gametocytes 
after entering a corpuscle are spherical with a distinct nucleus 
and without the vacuole typical of ring forms, while those 
destined to give rise to female gametocytes stretch across the 
corpuscle in the form of an elongate bar. 

Knowles and Das-Gupta (1931) observed scanty phago- 
cytosis in a case of infection with L. malarias, and Chopra, 
Das-Gupta, and Sen (1932) observed heavy phagocytosis of 
parasites by the polymorphonuclear and large hyahne leuco- 
cytes in another case. Stott (1933) has reviewed the previous 
hterature on the phagocytosis of malarial parasites and 
recorded his own observations in two cases. In severe 
malignant tertian infection approximately 50 per cent, of 
red blood-cells may be infected, and in such cases merozoite 
formation may be seen in the peripheral circulation. Large 
mononuclear cells may show their phagocytic power, even in 
the peripheral circulation of severe cases, by the presence of 
(1) pigment, (2) rings, (3) red cells, normal or dehsemoglobi- 
nized, and non-infected or infected with sporulating or, less 
commonly, with other forms of malarial parasites. 

MissiroH (1933), working with Culex fatigans infected with 
bird malaria, and Knowles and Basu (1935) with Anopheles 
stephensi infected with Laverania malariee and Plasmodium 
vivax, have observed division of the nucleus of the sporozoites 
into two, three or more parts. The sporozoite is then said to 
divide into a corresponding number of small rounded bodies 
consisting almost entirely of chromatin with scarcely any 
visible cytoplasm. These apply themselves to the surface 
of the red corpuscles and presumably grow into the usual ring- 
trophozoite forms. If these observations are confirmed it 
would mean that the sporozoite is not the end of the sporogony 
cycle, but that occasionally, at any rate, it breaks up into 
smaller bodies which, after entering the corpuscles, grow into 

ChaouHtch (1936) has recently observed that asexual 
multiphcation can also take place by binary fission of the 

Alessandrini (1933) reported a new method of reproduction 
for L. malarise, i. e., simple amitotic division, occurring both 
in man and mosquito, but only when the resistance of the host 
is distinctly lowered by infection, adverse environmental 
influences, etc. This amitotic reproduction results in a more 
rapid and virulent multiplication of the parasite, accounting 
for the occasional pernicious character of the disease. Hingst 
(1934) has also adduced evidence to show that in some indi- 


viduals of L. malar isR repeated direct division and formation 
of two daughter organisms of equal size occurs, and according 
to him this explains the multiple infection of the red blood- 
corpuscles so characteristic of the species. He believes 
that schizogony maj'- follow direct division. 

Boyd (1935) has discussed the comparative morphology 
of the sporozoites of the human malarial parasites. The 
form with dissimilar extremities appears most frequently in 
P. vivax, while in L. malarise both ends tend to be pointed. 
The sporozoites of L. malaria are the finest, those of 
P. malaria the coarsest. 

Cultivation. — Bass and Johns (1912) were the first to 
successfully cultivate this species outside the body, and to 
observe three generations of schizogony in these cultures. 
J. G. Thomson and T>. Thomson (1913) introduced a reliable 
modification of Bass and Johns's technique. Row (1917) 
successfully cultivated the malarial parasites in blood drawn 
from the finger, and pubHshed his observation on the forms 
observed in culture of Plasmodium falciparum and Laverania 
prsRCox (synonyms of Grassi and Feletti's two subspecies of 
Laverania (Hsemamoeba) malarise). In the former he noted 
the small size of the infected corpuscle and the formation of 
six or less merozoites : in the latter the mature schizont 
formed four or less merozoites and occupied the whole of the 
infected corpuscle, which was much larger than the normal one. 
The schizogony cycle completed itself in twenty-four hours, 
and there was a marked tendency on the part of the developing 
parasite to agglutinate in larger or smaller masses. Later 
workers do not regard the two subspecies as distinct. Sinton 
(1922) described the culture of malarial parasites from the 
finger-blood in specially constructed glass tubes. The details 
of some of these methods will be found in the part deaHng 
with practical methods. 

" Plasmodium tenue." — Stephens (1914), under the name of 
Plasmodium tenue, described a malarial parasite seen by him 
in a single blood-film sent to him from the Central Provinces. 
Only young forms were present, and were distinguished by 
their marked amoeboid form, large size of the nuclear chromatin, 
and markedly irregular shape. Balfour, Andrew, and Wenyon 
(1914) pointed out that such forms were not uncommon in 
Laverania malarias, and concluded that P. tenue was not a dis- 
tinct species. Knowles (1923 and 1926) expressed a similar 
opinion. Sinton (1922 b) found these forms first in five 
cases of malaria in the Central Provinces, and later in other 
cases at Lahore, and made out a strong case for the vahdity of 
P. tenue as a distinct species. Christophers (1925), in his 
study of malaria in Singhbhum, notes that the parasites 
encountered in that district resembled P. tenue rather than 


Laverania malariee. Callanan (1926) found " tenue " forms 
in blood-films containing Laverania malarise, and noted that 
the " tenue " forms tended to preponderate in certain portions 
of the film, whereas ring forms showed a predilection for others. 
He concluded that near the edge of the blood-film, where the 
red corpuscles are evenly distributed and lie singly, pressure 
in spreading the film causes the deHcate hair-like rings of 
Laverania malarise to become distorted into band-like " tenue " 
forms. So the case for P. tenue as a distinct species is by 
no means proved. 

Habitat. — Blood of man, in almost all parts of India, and 
the body of the following species of mosquitoes : — Anopheles 
annularis Ya,n der Wulp, A. culicifacies Giles {=A.fuliginosus), 
A. fluviatilis James {=A. listoni Liston), A. maculatus Theo., 
A. minimus Theo., A. philippinensis hudl., A.stephensi Liston, 
A. sundaicus Rodenwaldt, and A. varuna Iyengar. For the 
locahties in India, Burma, and Ceylon where Laverania 
malarise infection is known to occur, the map reproduced 
from Knowles, Senior White, and Das-Gupta (1930) at the 
end of the description of malarial parasites of man (p. 279) 
may be consulted, and for the relative importance of various 
species of Anopheles in the transmission of malaria in different 
parts, the useful summaries taken from Covell (1931) and 
Christophers (1933) may be referred to. 

Genus PLASMODIUM Marchiafava & Celli, 1885. 

Malarial parasites, Laveran, 1880 a, p. 158; 1881, pp. 627-30; 

1882, p. 737. 
Oscillaria (part), Laveran, 1883, p. 113. 
Plasmodium, Marchiafava & Celli, 1885, p. 791 ; Golgi, 1889, 

p. 173. 
Laverania -{-Hmmamceba, Laveran, 1890, p. 374 ; Grassi & Feletti, 

1892, p. 10. 
Hsemamceba, Labbe, 1894, p. 170. 
Plasmodium, Labbe, 1899, pp. 80-2. 
Heemomenas, Ross, 1899, pp. 322-4. 

Hcemamceba, Ross, 1899, pp. 322-4, 439 ; 1900, pp. 522-7. 
Plasm,odium, Liihe, 1900, pp. 367-84, pp. 436-60 ; Neveu-Lemaire, 

1900, p. 9 ; Minchin, 1903, pp. 239, 265, 267. 
Heemamceba (part), Laveran, 1907, pp. 110-60. 
Plasmodium, Minchin, 1912, p. 357 ; Row, 1917, p. 392 ; Castellani 

& Chalmers, 1919, pp. 504-16; Miihlens, 1921, pp. 1495-502; 

Thomson & Woodcock, 1922, pp. 1517-30 ; Hegner& Taliaferro, 

1924, pp. 317-40 ; Craig, 1926, pp. 362-470 ; Wenyon, 1926, 

pp. 908-85 ; Hehir, 1927, pp. 164-6 ; Knowles, 1928, pp. 382- 

410; Row, 1929, pp. 1120-5; Reichenow, 1929, pp. 998-1009; 

Kudo, 1931, pp. 286-7 ; Calkins, 1933, pp. 406-10, 566 ; Brumpt, 

1936, pp. 424-38. 

Trophozoites ring-shaped, amoeboid. Schizogony in the 
peripheral blood. Schizonts, compact or irregular, may 
completely fill the infected red blood-corpuscle, which may or 


may not be altered in size. Hsemozoin pigment present. 
Gametocytes more or less resemble the schizonts in form ; 
in the fully grown condition are usually circular discs which 
more or less completely fill the infected red blood- corpuscle. 
Parasites of man and other mammals. 

Remarks. — Coatney and Roudabush (1936) have given 
a Hst of all the known species of the genus Plasmodium together 
with the hosts from which each was described. 

232. Plasmodium vivax (Grassi & Feletti). (Fig. 123 (PI. I.)). 

Plasmodiutn var. tertiana, Golgi, 1889, p. 173. 

Plasmodium malariee var. tertianse, Celli & San Felice, 1891. 

Hcemamceba febris tertiana, Marchiafava & Bignami, 1891. 

Hsemamoeba vivax, Grassi & Feletti, 1892, p. 10. 

Hgatnamoeba laverani var. tertiana, Labbe, 1894, p. 170, pi. ix. 

Plasmodium malarise tertianum, Labbe, 1899, p. 82, fig. 147 a. 

Hcemamosba malarise var. magna, Laveran, 1900. 

Plasmodium vivax, Liihe, 1900, p. 460 ; Neveu-Lemaire, 1900, 
p. 9, pi. i, fig. 2. 

Hsemamoeba malarix var. tertianse, Laveran, 1901. 

Plasm,odium vivax, Schaudinn, 1902, pp. 169-250, pis. iv-vi ; 
Minchin, 1903, pp. 243-54, 267, 332, 351 ; Doflein, 1909, 
pp. 670-3, figs. 612-15 ; Minchin, 1912, pp. 137, 358, 359, 360, 
figs. 72, 156. 
■^Plasmodium vivax. Row, 1917, p. 392, pis. xx— xxii. 

Plasmodium, vivax,Cast6\\&-n.\ & Chalmers, 1919,pp. 510-12, figs. 166, 
169 ; pi. i, figs. 16-8 6; Miihlens, 1921, pp. 1495-9 ; pi. xxx, 
figs. 1-36; pl.xxxi,figs. 1-8; pl.xxxii, figs. 1-15 ; pi. xxxiii,fig. 4; 
Thomson & Woodcock, 1922, pp. 1517-29, pi. Ixiii, figs. 1-37 ; 
pi. Ixiv, figs. 38-47, 50-7 ; text-figs. 540-4 ; Hegner & Talia- 
ferro, 1924, pp. 317-26, 329-30 ; pi. i, fig. 124 ; Craig, 1926, 
pp. 393-413, figs. 67-70 ; Wenvon, 1926, pp. 925-34, pi. xii, 
figs. 1-35 ; fig. 400 ; Hehir, 1927, pp. 164-6, pi. x, figs. 1-37 ; 
pi. xi, figs. 38-47, 50-7 ; text-fig. 581. 
■fPlasrnodium vivax, Knowles, 1927, pp. 7-11, pi. i, figs. 1-31 ; 1928, 
pp. 383-9, pi. xi. 

Plasmodium vivax, Reichenow, 1929, pp. 998-1004, fig. 975, f-l, 
figs. 976-86 ; Kudo, 1931, p. 286, figs. 120, 121, a-g ; Calkins, 
1933,p.238,fig. 124,^, B; pp. 406-10, pi. i, figs. 1-6"; Brumpt, 
1936, pp. 432-8, figs. 190-2; Coatney & Roudabush, 1936, 
p. 340. 

The Cycle in Man. — Sporozoites are long and slender, 
10-12 ju- by 1-2^, and are introduced into the blood through 
the bite of an infected mosquito. After entering the corpuscle 
the sporozoite contracts to form a disc of cytoplasm with 
a single nucleus, and soon develops a vacuole, thus resembling 
a signet-ring. The rings are comparatively large, not as a rule 
at the edge of a red blood-corpuscle, and usually not more than 
one in each corpuscle. A ring occupies about one-third the 
diameter of the corpuscle. In blood-films of heavy infections 
there may also be young forms at the edge of the red corpuscles 
(marginal forms). They may be seen in stained specimens, 
either as little streaks of blue cytoplasm with a red nucleus 

272 spoRozoA. 

(flattened marginal forms) or disc-shaped bodies with a large 
central vacuole and with an arched appearance (bridge forms). 
In living blood, in the warm stage, the trophozoite may be 
seen actively throwing out and withdrawing pseudopodia 
within the corpuscle : this special activity earned the name 
P. vivax for the species. After further growth the trophozoite 
has a more irregular shape (amoeboid form) or the ring 
form continues, the vacuole becomes larger, and a few yellow 
or hght brown pigment granules are deposited in the cyto- 
plasm. Growth is accompanied by three distinct changes 
in the red blood- corpuscle ; it becomes larger, shghtly paler, 
and shows on its surface a number of fine granules, which 
stain bright pink by Romanowsky stains. These granules 
are known as Schiiffner's dots : they are not visible in poorly 
stained films. The schizont has the form of a circular plate 
of cytoplasm, which almost completely fills the enlarged red 
blood- corpuscle . Nuclear multipHcation takes place, producing 
twelve to twenty-four nuclei. The cytoplasm gradually 
segments into as many portions as the nuclei, and the merozoites 
separate, leaving the pigment in the residual cytoplasm. 
The merozoites are liberated into the plasma by the bursting 
of the corpuscle and enter other corpuscles. The majority 
of parasites complete schizogony at or near the critical period 
of forty-eight hours, though some may do so before or after. 
Sometimes double infections take place, sporozoites having 
been introduced on two successive nights. The schizonts 
of one batch will attain maturity one day and those of the 
other twenty-four hours later. Thus there will be daily attacks 
of fever instead of every other day. The initial attack of 
fever takes place about ten to twelve days after the intro- 
duction of the sporozoites by a mosquito. 

After schizogony has taken place a number of times some 
of the merozoites develop into gametocytes. This phase 
takes place mostly in the blood-vessels of the spleen or bone- 
marrow, as the red blood-corpuscles containing the developing 
gametoc3^es are held up in these organs. The gametocytes 
attain maturity in about ninety-six hours, and the effect 
produced on the red blood-corpuscles is the same as in the 
case of the schizonts. The gametocytes are circular in outhne, 
more regularly circular than the schizont ; each contains 
a larger number of pigment granules, more or less uniformly 
distributed in the cytoplasm, and a single nucleus. The 
male gametocyte has hyaline cytoplasm which stains a pale 
blue, and the nucleus is larger and contains fine chromatin 
granules. The female gametocjrte has denser cytoplasm, 
which stains more deeply blue, has a smaller nucleus, with 
a single karyosome or a group of granules. Both contain 
yellowish -brown pigment granules distributed in the cytoplasm. 


The Cycle in the Mosquito. — ^The development in the mos- 
quito is on the same Hnes as in Laverania malarise. As the 
gametocytes of P. vivax are the largest, the zygotes and 
oocysts of this species are also probably larger than in the 
other species. The pigment produced in this species is of 
a hght brown colour, while that in the other two species of 
human malarial parasites is dark brown or black : thus 
zygotes and oocysts with light brown pigment will be those 
of P. vivax. 

Remarks. — Grassi (1900) and Schaudinn (1902 a) described 
certain changes in the female gametocyte which they inter- 
preted as parthenogenesis. J. D. Thomson (1917) and Wenyon 
(1926) have refuted the theory and explained that the appear- 
ances observed may be due to a red corpuscle being infected 
with two or more parasites, which may both be schizonts, 
or a schizont and a gametocyte, and thus combinations of all 
sorts of appearances may result. 

Chalmers and Archibald (1920) described what they call 
the " tenue " phase of P. vivax, having the same relation to 
this species as P. tenue Stephens has to Laverania malarias. 

James, Nicol, and Shute (1933) gave a description of the 
life-cycle of Plasmodium ovale Stephens, with different 
characters to separate the species from the other malarial 
parasites. According to them the parasite is identical with 
that described by Craig in 1900 and 1914, but is distinct from 
P. camarense Emin. Giovannola (1935) beheves that P. ovale 
is a modification of P. vivax after a long residence in the 
human host, and cites historical, cHnical, and morphological 
evidence to support his claim. 

Habitat. — Blood of man in almost all parts of India {vide 
map on p. 279), and in the body of several species of Anopheles, 
such as A. annularis Van der Wulp, A. culicifacies Giles, 
A. fluviatilis James (=.4. listoni Liston), A. maculatus Theo., 
A. minimus Theo., ^. philippinensis Ludl., A. stephensi Liston, 
A. sundaicus Rodenwaldt, and A. varuna Iyengar. 

233. Plasmodium malariae (Laveran). (Fig. 124 (PL I).) 

Malarial parasites, Laveran, 1880 a, p. 158 ; 1881, pp. 627-30 ; 

1882, p. 737. 
Oscillaria malar iee (part), Laveran, 1883, p. 113. 
Plasmodium var. quartana, Goigi, 1885 ; 1886 ; 1890. 
Amoeba malarise febris quartanee, Celli, 1891. 
Plasmodium, malarise quartanee, Celli & San Felice, 1891. 
Hcemam,ceba febris qu^artanse, Marchiafava & Bignami, 1891. 
Hsemammba malarise, Grassi & Feletti, 1892, p. 10. 
Hsemamceba laverani var. quartanarum, Labbe, 1894, p. 170. 
^Hssmam^cBba malarise, Ross, 1898, pp. 133—6. 
Plasmodium malarise quartanum, Labbe, 1899, p. 82, fig. 147 b. 
Plasmodium malarise, Liihe, 1900, p. 460 ; Neveu-Lemaire, 1900, 
p. 8, pi. i, fig. 1. 


Plasmodium malarias, Minchin, 1903, pp. 243, 267 ; 1912, p. 358. 
fThe Quartan Parasite, Row, 1917, p. 392, pis. xviii, xix. 

Plasmodium malarise, Castellani & Chalmers, 1919, pp. 512-13 ; 
Muhlens, 1921, pp. 1499-502, pi. xxxi, figs. 9-19 ; pi. xxxii, 
figs. 16-25 ; pi. xxxiii, figs. 1, 3 ; Hegner & Taliaferro, 1924, 
pp. 326-7, 330, 331, pi. ii, figs. 1-8 ; Craig, 1926, pp. 419-31, 
figs. 72, 73 ; Wenyon, 1926, pp. 942-4, pi. xiii, figs. 1-15 ; 
Hehir, 1927, pp. 162-4, pi. ix, figs. 1-3 ; pi. xi, figs. 58-75. 
■^Plasmodium, maJarisR, Knowles, 1927, pp. 12-14, pi. ii, figs. 1-22 ; 
1928, pp. 389-92, pi. xii. 

Plasmodium malarise, Reichenow, 1929, pp. 1004-6, fig. 975, m-p, 
987-90; Kudo, 1931, p. 287, fig. 121, o-m ; Calkins, 1933, 
pp. 238, 406-10, fig. 124 ; pi. i, figs. 7-12 ; Brumpt, 1936, 
pp. 424-30, pi. ii, figs. 19-36; Coatney & Roudabusb, 1936, 
p. 339. 

The Cycle in Man. — The cycle in man is very similar to 
that of Laverania malarias and Plasmodium vivax. " Rings " 
are of the same size as of P. vivax and have a diameter of about 
one-third to a half that of the red blood-corpuscle. Cytoplasm 
is denser and stains more deeply, and shows little amoeboid 
activity during growth. Pigment granules are coarse and of 
a darker brown colour. The organism often shows a tendency 
to be stretched as a band across the diameter of the corpuscle. 
These " band forms " are more frequently seen in this species. 
The band may be narrow or almost as broad as long. The 
infected red corpuscle is no larger, and may even be smaller 
than the normal corpuscle, and may also be deeper in colour 
in stained films ; and as a rule no Schiiffner's or Maurer's dots 
are found. Schizont reaches its full size in about sixty 
hours and schizogony occurs every seventy-two hours. The 
fully grown schizont is circular in outhne, and almost com- 
pletely fills the corpuscle, the dark pigment granules forming 
a central mass. Schizogony results in the production of six 
to twelve merozoites that are arranged ia a single ring or 
" rosette." All phases of schizogony take place in the peri- 
pheral blood. The gametocytes are oval or spherical in outHne, 
and completely fill the red blood- corpuscles. The distinguishing 
characteristics of microgametocytes and macrogametocytes 
are similar to those in P. vivax, the female staining more deeply 
blue and having a more compact nucleus. The pigment 
granules are scattered irregularly in the cytoplasm. 

The Cycle in the Mosquito. — The development in the Ano- 
phehne mosquito is very similar to that of P. vivax, but the 
various stages can be distinguished from those of P. vivax 
by the pigment being dark brown or black. In this respect 
they resemble those of Laverania malarias. 

Remarks. — Two " rings " or partially developed tropho- 
zoites may sometimes be found in the same corpuscle, but are 
less common than in P. vivax infections. Two mature 
gametocytes in an enlarged cell, as also a gametocyte and a 
schizont, have also been found to occur in the same corpuscle. 


Double and triple infections sometimes occur owing to 
inoculations by mosquitoes on different days. Each group 
of parasites undergoes schizogony independently, thus attacks 
of fever occur in cases of single infection every seventy- two 
hours, that is, on the fourth day (hence quartan). Double 
infection may cause the attacks to appear on two successive 
days, and then again on two successive days after an interval 
of twenty- four hours. In cases of triple infection attacks of 
fever take place every twenty-four hours. 

Mixed infection of more than one species may also occur. 
Very often P. vivax and Laverania malaria are found in the 
same blood-film, as both these species are of commoner 
occurrence than P. malarise. But the latter may also be met 
with in association with either of the others, and occasionally 
all three have been seen in the same blood-film. The diagnosis 
of these mixed infections will depend upon the finding of 
undoubted forms of each species. 

Habitat. — Blood of man in almost all parts of India {vide 
map on p. 279) and in the body of several species of Anopheles, 
such as A. annularis Van der Wulp, A. culicifacies Giles, 
A. fluviatilis James {^=A. listoni Liston), A. maculatus Theo., 
A. minimus Theo., A. philippinensis Ludl., A. stephensi 
Liston, A. sundaicus Rodenwaldt, and A. varuna Iyengar. 

Distribution of the Human Malarial Parasites. 
The climatic features of the different part of India being 
so diverse, greatest diversity of distribution of species of 
malarial parasites is found in the country. Knowles, Senior 
White, and Das-Gupta (1930) have come to the following 
general conclusions on the basis of the available data : — 

(1) In the north-west of India P. vivax tends to predominate 
during the spring and early summer and L. malarise in the 
late autumn. P. malarise is quite unimportant. 

(2) The importance of P. malarise becomes greater and 
greater as one passes from the N.W. Frontier Province to the 
east and south-east. In brief we may trace two chains of 
increasing quartan prevalence, as follows : — 

(a) Across Northern India the figures run : 

N.W.F.P., 0-3 per cent. ; Punjab, 1 ; Delhi Province, 
1-2 ; U.P., 4-6 ; Bengal, 12-3 ; Assam, 9-9 ; Burma, 

{b) Down the west coast of India : 

Sukkur and Sind, 15-2 per cent. ; Bombay City, 6-6 ; 
Goa, 9 ; Kanara, 2-1 ; Mysore State, 15-7 ; Coorg, 
24 ; Ceylon, 33 (in some places up to 71 per cent.). 
Other hilly areas where the incidence is high are the 
Central Provinces, 19 per cent., and the Madras Agency 
Tracts, 41. 






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(3) Taking all-India generally, P. vtvax is the predominant 
species, but this only appHes to the spring and early summer 
months. In autumn and early winter L. malaria usually 
becomes predominant. 

(4) Benign tertian {P. vivax) and quartan malaria {P. malaria) 
are almost equally common in Ceylon, although in certain parts 
of the island — and especially in the hilly districts — quartan 
malaria predominates. The figure for maHgnant tertian 
{L. malar ise) is extremely low throughout the island. 

Taking the figures for India as a whole, it is clear that 
benign tertian and maHgnant tertian infections share the main 
incidence of malaria in India about equally between them, 
while quartan malaria is responsible for less than 7 per cent. 
The relative distribution of the different species in various 
localities will be seen from the map reproduced from Knowles, 
Senior White, and Das-Gupta on p. 279. 

Distribution of Mosquito-carriers of Malaria. 

Covell (1931) has summarized the available information 
concerning the mosquito carriers of malaria in India, Burma, 
and Ceylon as follows : — 

" The chief malaria carriers in Northern and Peninsular 
India are A . culicifacies, A . stephensi, and A . listonii. A . culici- 
facies is the most important vector in rural areas, whilst 
A. stephensi is notorious as being the great urban malaria 
carrier of India, it being the only malaria- carrying species 
capable of adapting itself to the conditions obtaining in cities. 

"A. maculatus and its variety willmori are considered to be 
important carriers in submontane areas, chiefly on epidemio- 
logical grounds. A. maculatus has been found naturally 
infected (gut only) by Feegrade (1927 a) in Burma. 

' ' In Eastern India A . minimus is probably the most important 
carrier. Ramsay (1930) has shown this species to be the 
principal vector in the Cachar District of Assam. Iyengar 
(1927) considers A. minimus var. varuna to be the chief 
carrier in Lower Bengal. Sur and Sur (1929), as the 
result of their dissections under natural conditions, conclude 
that A. philippinensis plays an important part in malaria 
transmission in Bengal. This species has also been found 
naturally infected in Burma (Feegrade, 1926). A. fuliginosus 
has also been found infected in nature in Bengal and in Madras, 
but this species is not generally considered to play an important 
part in transmission. 

"A. ludlowii is an important carrier in the Andamans and 
on the coasts of Burma and Bengal. In the Andamans it is 
considered to be the only vector of any practical importance. 





" Various other species have been suspected to be carriers 
of malaria in India, but there is no direct evidence that they 
play any important part in the transmission of the disease. 

" In Ceylon Carter (1927) considers that A. culicifacies 
and A. listonii are the chief carriers, whilst the position of 
A. maculatus is uncertain. The last-named species, although 
prevalent in certain districts where malaria is severely endemic, 
is relatively more abundant at somewhat higher elevations 
(1500 feet and over), where the incidence of the disease is low, 
Ibut where the factors of temperature and atmospheric humidity 
are favourable for transmission. Senior White (1920), 
however, is of opinion that this species is ' the malaria carrier 
par excellence of the Ceylon hill country,' on epidemiological 

Christophers (1933) has summarized the available data 
as follows : — 

"1. Important carriers wherever found : 

A. culicifacies ; A. fiuviatilis ; A. stephensi ; 
A. sundaicus ; A. minimus. 

"2. Less important, but proved carriers in some areas : — 
A. varuna ; A. philippinensis . 

" 3. Species that are important carriers in other countries, 
but of too limited distribution to be important in 
India : 

A. superpictus ; A. multicolor. 

*' 4. Species that have been found infected in nature or 
experimentally within or without the area, but which 
are probably not of importance as carriers : 

a. Found infected in nature in Indian area : 

A. maculatus ; A. maculatus var. willm,ori ; 
A. fuliginosus ; A. pulcherrimus ; A. maculi- 
palpis ; A. pallidus ; A. ramsayi ; A. vagus. 

b. Infected experimentally only in Indian area : 

A. theohaldi ; A. suhpictus ; A. turkhudi. 

c. Found infected in nature outside Indian area only : 

A. hyrcanus ; A. harhirostris ; A. karwari ; 
A. leucosphyrus ; A. sergenti ; A. umbrosus ; 
A. tassellatus ; A. aconitus ; A. kochi. ■ 

" The extremely common species A. suhpictus and A. vagus 
appear to have little or no relation to the incidence of 

Iyengar (1934) found five species of Anopheles, viz., 
A. jeyporiensis var. candidiensis Koidz, A. varuna Iyengar, 
A. fiuviatilis James, A. listoni (Liston), and A. culicifacies 
Giles, infected in nature in Travancore. 


234. Plasmodium cynomolgi Mayer. (Fig. 125 (PI. II).) 

Plasmodium cynomolgi, Mayer, 1908, pp. 314-22, pi. xxi ; Flu, 
1908, pp. 323-30, pi. xxii ; Blanchard & Langeron, 1913, 
pp. 529-42, pis. viii-ix ; pp. 599-607, pi. x ; Castellani & Chal- 
mers, 1919, p. 515. 
■fPlasmodium cynomolgi (?), Donovan, 1920, p. 721. 
Plasmodium cynomolgi, Miihlens, 1921, p. 1612-14, fig. 32 ; Wen- 
yon, 1926, p. 969. 
Plasmodium inui, Wenyon, 1926, p. 971 ; Knowles, 1928, p. 439 ; 

Reichenow, 1929, p. 1007, fig. 993. 
Plasm,odium cynomolgi, Reichenow, 1929, pp. 1007—8, fig. 994. 
Plasmodium inui (?),' Green, 1931, pp. 649-50 ; 1932, pp. 455-77, 
7 figs. 
■\Plasmodium kochi (?) (part), Napier & Campbell, 1932, pp. 246-9. 
^Plasmodium sp. (part), Knowles & Das-Gupta, 1932, pp. 300-20, 

7 pis. & 6 charts ; Sinton & MuUigan, 1932, p. 324. 
■fPlasmodium inui var. cynomolgi, Sinton & Mulligan, 1932 a, 

pp. 396-405. 
'\Plasmodium cynomolgi, Sinton, 1934 a, pp. 48—50 ; 1934 6, 
pp. 392, 399-400 ; Mulligan, 1935, pp. 288-300, pi. iv, figs. 1-35. 
Plasmodium cynomolgi, Coatney & Roudabush, 1936, p. 339. 

Young rings, 2-5 ju, in diameter, round, or oval, with promi- 
nent chromatin dot and well- developed vacuole. Two or 
three rings sometimes seen in same red corpuscle ; infested 
corpuscle usually unaltered ; no stippling. Growing forms 
often show marked amoeboid movement, resembb'ng that seen 
in P. vivax. Pigment scanty, appears later, and is darker and 
coarser than in P. inui ; vacuolation, especially in large 
forms, not marked ; infected red corpuscles appreciably 
enlarged and pale ; stippling resembling Schiiffner's dots, 
conspicuous. Schizonts common in peripheral blood ; mature 
schizonts with eight to sixteen merozoites irregularly scattered ; 
pigment in small dense clump ; infected red corpuscles with 
marked pallor and prominent stippling. A minute accessory 
chromatin dot frequently seen in merozoites and young rings. 
Schizogony cycle takes forty-eight hours to complete. Game- 
tocytes intra-corpuscular or free, larger than the normal red 
corpuscle. Female gametocytes round or oval ; cytoplasm 
deep blue, chromatin small, compact, and eccentric ; pigment 
not very abundant, and granules darker and coarser than in 
P. inui. Male gametocytes smaller, cytoplasm staining 
less deeply, chromatin central or peripheral, larger and less 
deeply staining than in the female gametocytes. Sporogony 
cycle in mosquitoes. 

Remarks. — P. inui was described by Halberstadter and 
Prowazek (1907) from monkeys {Macaco) from Borneo. 
Mayer (1908) described P. cynomolgi, which was generally 
considered as identical with P. inui. He (1908) could obtain 
no development in Culex pipiens nor in Aedes (Stegomyia) 
segypti (Linn.) [^ Aedes argenteus] ; he observed small oocysts 
in Anopheles maculipennis, but could not follow the complete 


development. Leger and Bouilliez (1913) inoculated the 
parasite into a number of monkeys, including Cercopithecus sp. 
(the common host of P. kochi), with the result that many of 
them died of heavy infections. Quinine, even in large doses, 
appeared to have no influence. Donovan (1920) examined 
a large number of specimens of Silenus sinicus and Pygerythrus 
priamus {^=Preshytis j^riamus) in the Nilgiri Hills, South India, 
but with negative results. Subsequently he found a Plasmo- 
dium, in a blood-film from a specimen of S. sinicus from the 
same region, which he considered as morphologically identical 
with that found in 8. irus {=M. cynomolgus). He also 
mentions having seen Plasmodium in several other apes 
and monkeys, but does not give any description or cite 

Francliini (1927) found in Macacus cyoiomolgus a parasite 
which showed many resemblances with P. malariss. Macfie 

(1928) found P. inui in a young baboon in Africa. Reichenow 

(1929) observed that P. inui does not show enlargement 
of the corpuscles and Schiiffner's dots, whilst P. cynomolgi 
shows these, but stilJ came to the conclusion that these forms 
are probably identical. 

Sinton and Mulligan (1932), in a specimen of Silenus irus, 
obtained an infection which later work showed was a mixed 
one. They succeeded in isolating in S. rhesus, by blood 
inoculation from the same naturally infected host, two mor- 
phologically distinct types of Plasmodium, which they regard 
as P. inui var. cynomolgi and P. knowlesi. They obtained 
the sporogony cycle in a number of mosquitoes, and succeeded 
in transmitting the infection to a healthy specimen of Silenus 
rhesus by the bites of infected A. annularis Van der Wulp. 
Napier and Campbell (1932) and KJnowles and Das- Gupta 
(1932) also recorded a malarial infection in 8. irus, said to 
have been imported from Singapore. Sinton and MulHgan 
(1932 a) gave a critical review of the literature relating to 
malarial parasites in lower monkeys, and came to the con- 
clusion that the majority of species of Plasmodium described 
from the lower monkeys fall into two main divisions, those 
found in the African monkeys {P. kochi group) and those found 
in the Oriental ones (P. inui group). According to them the 
latter include P. inui Halb. & Prowazek {sensu restr.), P. inui 
var. cynomolgi Mayer, P. inui var. gonderi, and P. knowlesi, 
and the various previous records were arranged according 
to this scheme. Later on, Sinton (1934 a, 1934 b) and Mulligan 
(1935) have come to the definite conclusion that P. cynomolgi 
is a distinct species and not a variety of P. inui, as the duration 
of schizogony in the former is forty-eight hours and in the 
latter seventy- two hours. 


Habitat. — Blood of Silenus sinicus (Linn.) Nilgiris ; blood of 
Silenus irus (Cuv.), said to have been imported from Singapore, 
and in inoculation infections in S. irus (Cuv.), S. rhesus 
(Audeb.), and S. sinicus (Linn.) ; sporozoites in the salivary- 
glands of Anopheles annularis Van der Wulp {=A. fuliginosus 
Giles), A. splendidus Koidzumi (=^. maculipalpis James & 
Liston), A. maculatus Theobald, and A. culicifacies Giles : 
Bengal, Calcutta ; Punjab, Kasauli. 

235. Plasmodium inui Halberstadter & Prowazek. (Fig. 126 
(PI. II).) 

Plasmodiwn inui, Halberstadter & Prowazek, 1907, pp. 37-43, 
pi. vi; Castellani & Chalmers, 1919, p. 515; Miihlens, 1921, 
p. 1612 ; Wenyon, 1926, pp. 971-2, pi. xv, figs. 8-14 ; Knowles, 
1928, p. 439 ; Reichenow, 1929, pp. 1007-8, fig. 993. 

Plasmodium inui, sens, restr., Sinton & Mulligan, 1933, pp. 389- 

^Plasmodium inui, Sinton, 1934 h, pp. 393-410, pis. iii, iv ; Mulligan, 
1935, p. 310. 

Plasmodium inui, Coatney & Roudabush, 1936, p. 339. 

Young rings about one-fifth to one-fourth diameter of in- 
fected red ceU ; marked vacuolated appearance, amoeboid 
movement of lobose nature. Older trophozoites more rounded, 
mature forms do not fill infected red cell. Mature schizonts 
with sixteen merozoites which often form a rosette. Chromatin 
relatively large, prominent, and usually excentric at all stages ; 
may be divided into two equal or unequal masses in young 
forms. Pigment yellow to brown, becoming darker with age ; 
appears early ; fine and abundant with peripheral distribution. 
Infected red cells shghtly enlarged with older forms ; stipphng 
less conspicuous, scantier than with P. cynomolgi. Duration 
of schizogony cycle seventy- two hours. Gametocytes rounded, 
about the size of normal red cells, with yellowish-brown to 
brown pigment, scattered and abundant. Female gametocyte 
with cytoplasm stained deep blue with Romanowsky's stain, 
with a small chromatin mass, and with scattered and coarser 
pigment ; when mature, larger than normal red corpuscle, 
Male gametocyte with large loose chromatin mass and 
abundant scattered pigment, lighter brown than in the 

Remarks. — Sinton (1934 h) has come to the conclusion that the 
duration of schizogony is seventy-two hours, and not forty- 
eight hours as reported by MuUigan (1935), to which paper he 
had access prior to pubHcation. 

Habitat. — Blood of Silenus irus (Cuv.) from Malaya, and in 
inoculation infections in 8. irus (Cuv.) and S. rhesus (Audeb.) : 
Punjab, KasauH. 


236. Plasmodium knowlesi Sinton & Mulligan. (Fig. 127 
(PI. II).) 

Plasmodium sp., Franchini, 1927, pp. 467-71, 2 figs. 
Plasmodium kochi (?) (part), Napier & Campbell, 1932, pp. 246-9. 
Plasmodium, sp. (part), Kjiowles & Das-Gupta, 1932, pp. 301-20, 

7 pis. & 6 charts ; Sinton & Mulligan, 1932 a, p. 324. 
■\Plasm,odium, knowlesi, Sinton & Mulligan, 1932 6, pp. 379-80, 

409-21, pi. V, figs. 1-35. 
Plasmodium knowlesi, Coatney & Roudabush, 1936, p. 339. 

Youngest rings closely resemble those of Laverania malarise 
and measure one-fourth to one-half of the infected corpuscle. 
Cytoplasm shows sHght but definite thickening on the side 
opposite the chromatin, and a well- developed vacuole. 
Chromatin prominent, and occurs as a single round, oval or 
elongate mass or in two (rarely three) smaller masses. One 
or, rarely, two minute accessory chromatin dots may be 
present. Older trophozoites rounded or only very shghtly 
amoeboid, vacuole very inconspicuous or absent. Mature 
schizonts of the same size, or smaller than the normal red 
corpuscles. Number of merozoites commonly about ten. 
Schizogony cycle takes twenty-four hours. Pigment appears 
early, is relatively abundant, and granules are fairly coarse, 
varying from greenish-brown to almost black. Infected 
red corpuscles not enlarged, but often show characteristic 
distortion, and stippling usually absent with ordinary stains. 
Oametocytes spherical, like those of P. malarise, about the 
size of normal red corpuscle, and with abundant coarse and 
dark pigment. Cytoplasm of the female gametocyte stains 
deep blue with Romanowsky's stain, and its chromatin is 
dense and compact, is usually situated peripherally, and 
frequently shows a more deeply staining inner area ; the 
pigment is scattered irregularly. The male gametocyte 
stains poorly, and its chromatin is large and diffuse, appearing 
to merge into the cytoplasm. Dark brown or almost black 
granules of pigment occur in both the female and the male 
gametocytes. Sporogony not known to take place in mos- 

Remarks. — Napier and Campbell (1932) found a Plasmodium 
in the blood of a specimen of Silenus irus in Calcutta, said 
to have been imported from Singapore. Knowles and Das- 
Gupta (1932) described it in greater detail after inoculating 
S. rhesus with the strain, in which host it multiphed enormously. 
Sinton and Mulligan (1932) came to the conclusion that the 
original infection in S. irus was a mixed one of Plasmodium, 
inui var. cynomolgi and of P. knowlesi, and by inoculating 
8. rhesu,s an almost pure infection with P. knowlesi was the 
result. A large number of specimens of Silenus rhesus 
from Northern India, examined by Knowles and Das-Gupta 
(1932) as well as by Sinton and Mulligan, have not shown 


any malarial infection, which suggests that this species of 
monkey rarely, if ever, suffers from natural infection with 
malaria in this region. 

Eaiowles and Das-Gupta (1932) were also successful in 
infecting a man with a Plasmodium from the lower monkeys. 

Habitat. — In natural infections of blood of Silenus irus 
(Cuv.) believed to be imported from Singapore, and experi- 
mental infections of S. rhesus (Audeb.) : Bengal, Calcutta. 

237. Plasmodium kochi Laveran. (Fig. 135.) 

Hsemamosba kochi, Laveran, 1899, p. 124. 
Plasmodium kochi, Leger & Boiiilliez, 1914, pp. 954—85. 
'fPlasmodium kochi, Castellani & Chalmers, 1913, pp. 387—8 ; 

1919, p. 515. 
Plasmodium kochi, Muhlens, 1919, pp. 1608-11, fig. 31. 
jPlasmodimn kochi, Donovan, 1920, p. 719. 

Plasmodium kochi, Wenyon, 1926, p. 971, pi. xv, figs. 22—8 ; 
Knowles, 1928, p. 439, fig. 102, 11-12 ; Reichenow, 1929, 
p. 1006, fig. 991. 
^Plasmodium, kochi, Sinton & Mulligan, 1933, p. 424. 
Plasm,odium, kochi, Coatney & Roudabush, 1936, p. 339. 

Resembles P. vivax. Ring-forms large. The young rings 
occupy a smaller proportion of the corpuscle than in P. vivax, 
but as growth proceeds resemblance becomes marked. The 
infected red blood -corpuscles become enlarged, parasite 
becomes irregular in shape, pigment granules are of a hght 
brown colour, and sometimes Schiiffner's dots may be found 

Fig. 135. — Plasm,odium, kochi (Laveran). A, ring-form ; B, schizont. 
(From Reichenow, after Gonda and Berenberg-Gossler.) 

to occur. Schizogony is completed in forty-eight hours. 
Schizont produces eight to fourteen merozoites, and bears 
a striking resemblance to that of P. vivax. Gametocytes 
are large round bodies which can be distinguished as male 
and female, as in P. vivax. 

Remarks. — The species is a common parasite of monkeys 
in tropical Africa, but shows little sign of pathogenicity in 
natural infections or in inoculated animals. Gonder and 
Rodenwaldt (1910) noted, however, that if splenectomy is 
previously performed infections are much more severe, 
temperature rises, and the parasites continue in the blood 
for many months. They were unable to inoculate the parasite 


into man. Further development could not be achieved in 
Anopheles maculipennis Meig. or in Aedes [Stegomyia) segypti 
(Linn.) [= Stegomyia fasciata]. Castellani and Chalmers (1913) 
stated that illness and death may occur among the monkeys 
in Ceylon, due to a malarial parasite which they refer to 
P. kochi. No description of the parasite is given, and according 
to Sinton and Mulhgan (1932) this report needs confirmation. 
Habitat. — Blood of monkeys : Ceylon ; Cercopithecus sp. ? : 

238. Plasmodium pitheci Halberstadter & Prowazek. (Fig. 

■^Plasmodium sp., Laveran, 1905 (as noted in Wenyon, 1926, p. 1363). 
Plasmodium pitheci, Halberstader & Prowazek, 1907, pp. 37-43, 

pi. vi; Shibayama, 1910, pp. 189-91, 1 pi. 
f Plasmodium pitheci, Donovan, 1920, p. 719. 
Plasmodium 2^it^ieci, Reichenow, 1920 c, pp. 207-16, 1 pi. ; 
Miihlens, 1921, pp. 1611-12. 
"^Plasmodium pitheci, Donovan (first recorded in Wenyon, 1926, 

p. 1363). 
Plasmodium pitheci, Wenyon, 1926, p. 972, pi. xv, figs. 29-35 ; 
Knowles, 1928, fig. 102, 13, 14 ; Reichenow, 1929, <^p. 1006-7, 
fig. 992 ; Coatney & Roudabush, 1936, p. 340. 

Young rings resemble those of Laverania malarias. FuUy 
developed trophozoite shows dark brown or black pigment. 
Schiiffner's dots present. Schizogony resembles that of 
P. vivaz. Gametocytes resemble those of P. malaria. 

Fig. 136. — Plasmodium pitheci Halb. & Prow. A, ring-form, double 
infection ; B, older ring ; C, schizont ; D, macrogamete ; 
E, microgametocyte. (From Reichenow, after Halber- 
stadter and Prowazek.) 

Remarks. — The parasite resembles P. inui, but is distinguished 
by its dark brown or black pigment. Shibayama (1910) states 
that Schiiffner's dots are not present, but Wenyon thinks 
that his staining was not sufficiently intense to show them. 
Dodd (1913) recorded an infection of an orang-utang with this 
parasite, which proved fatal. Reichenow considers it possible 
that what has been described as P. pitheci may have been 
one of the human parasites. 

Habitat. — Blood of orang-utang, Simia satyrus Desmarest : 
Asia (? exact locality — not given by Laveran) ; an Indian 
menagerie {Donovan). 



239. Plasmodium semnopitheci Knowles. (Fig. 137.) 

■^Plasmodium semnopitheci, Knowles, 1919, pp. 195-202, pis. xvi- 

Plasjnodium semnopitheci, Miihlens, 1921, pp. 1614—15. 
■^Plasmodium sp., Chimisso, 1922, pp. 38-54. 
Plasmodium, semnopitheci, Wenyon, 1926, pp. 972, 1363. 
■^Plasmodium semnopithecus, Scott, 1926, p. 237 ; Knowles, 1928, 
p. 439. 
Plasmodium semnopithecus, Reichenow, 1929, p. 1008 ; Sinton & 
Mulligan, 1933, pp. 382-6, 423-4; Coatney & Roudabush, 
1936, p. 340. 

Young trophozoite an almost non-pigmented ring. Growing 
trophozoites large flimsy rings with very large vacuole, very 
abundant pigment, little amoeboid movement, and single, 
oval or rod- shaped chromatin mass. Absence of segmenting 
forms in the peripheral blood. Decolorization of infected 
Ted corpuscles frequently observed, but stippHng never seen. 
Gametocytes rounded. Female gametocytes often free, and 
larger than normal red corpuscles ; chromatin small, deeply 
staining, oval or much elongated ; pigment very dark brown, 

A B C D 

Fig. 137. — Plasmodium semnopitheci Knowles. A, ring-form ; B, larger 
and more flimsy ring ; C, extracellular form impinging 
against a corpuscle ; D, gametocyte. (After Knowles.) 

very abundant, and in very fine granules or small dark collec- 
tions. Male gametocytes smaller than female gametocytes, 
and with very large pink nuclei. 

Remarks. — Knowles (1919) discovered this parasite during 
^n experiment conducted for the purpose of transmitting the 
mahgnant tertian parasite of man to a monkey. The shock 
of the operation and the injection of foreign blood were beheved 
to have stimulated a latent infection of a species peculiar 
-to the host, which suddenly became virulent and proved fatal. 
Nearly every corpuscle was found infected, and there were 
inumerable free forms, some of them undergoing schizogony 
whilst still extracellular. Wenyon (1926) thinks this due 
to intensity of infection and to examination after death : 
he considered the parasite to be very similar to P. inui, and 
Reichenow (1929) thought that it was probably identical with 
it. Sinton and Mulligan (1932 a) consider it noteworthy 
-that, except for Donovan's undescribed parasite, the only 



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natural malarial infections which have been reported from 
Indian monkeys, namely, by Klnowles (1919), by Chimisso 
(1922), and by Scott (1926), all appear to be due to P. semno- 

Habitat. — Blood of the entellus monkey or hanuman, 
Pygathrix entellus (Dufr.) {= Semnopithecus entellus) : Assam ; 
blood of Presbytes pileatus Blyth : Assam, in the Zoological 
Gardens, London. Also blood of Silenus rhesus (Audeb.), 
believed to be from India, examined in Italy. 

240. Plasmodium bubalis Sheather. (Fig. 138.) 

■fPlasmodium bubalis, Sheather, 1919 a, pp. 1-5, pis. i, ii ; 1919 b, 

p. 223. 
Ptasmodium bubalis, Muhlens, 1921, p. 1616. 
■^Plasmodium bubalis, Edwards, 1925, p. 50. 

Plasmodium bubalis, Wenyon, 1926, pp. 975-6, pi. xvi, figs. 13-17 ; 
Knowles, 1928, p. 442, fig. 102, 15, 16 ; Reichenow, 1929, p. 1009 ; 
Coatney & Roudabush, 1936, p. 338. 
^Plasmodium bubalis, Taylor, 1938, p. 42. 

Resembles P. malarias in many respects. Smallest parasites 
round, oval or pyriform ; pigment has appeared in parasites 
3/x in size. Larger parasites, 6^ in size, generally round, 

A B C D E 

Fig. 138. — Plasmodium bubalis Sheather. A, young trophozoite ; 
B, double infection with two trophozoites ; C, large 
trophozoite ; D, schizont ; E, merozoites escaping from the 
corpuscle. (After Sheather.) 

sometimes oval, and possess a central vacuole. Chromatin 
is peripheral, single or double, and the pigment in the cyto- 
plasm is scattered or in clumps. The larger parasites have 
a regular outHne and cause a shght enlargement of the infected 
red blood- corpuscle. Adult schizont completely fills the cor- 
puscle, produces seven to fourteen merozoites, and the pigment 
is aggregated in a single clump. Some of the large forms with 
a single nucleus are probably gametocytes, and many of them 
possess a vacuole. 

Remarks. — Sheather (1919) found the parasite in a buffalo, 
which died after inoculations made for immunization purposes, 
and noted that there was a fairly heavy infection, 1*6 percent, 
of the red corpuscles containing the parasite. Edwards 
(1925) noted the parasite again. 

Habitat. — Blood of buffalo. Bos bubalus Linn., and of ox, 
Bos indicus Linn. : United Provinces, Muktesar, Izatnagar. 

SPOK. tr 



241. Plasmodium canis Castellani & Chalmers. (Fig. 139.) 

■fPktsmodium canis, Castellani & Chalmers, 1910; 1919, p. 515, 

figs. 171, 172. 
Plasmodium canis, Muhlens, 1921, p. 1617 ; Wenyon, 1926, p. 976 ; 
Knowles, 1928, p. 442 ; Reichenow, 1929, p. 1009 ; Coatney & 
Roudabush, 1936, p. 338. 

Morphologically similar to P. vivax. Small round merozoite 
enters the red cell, and grows into a pigmented trophozoite, 
finally dividing into a number of merozoites. Schiiffner's 
dots are seen. Female gametocyte has a small rounded 
nucleus. Male gametocyte has a narrow elongated nucleus. 
Infected red cells are enlarged. 


Pig. 139. — Plasmodium, canis Castellani & Chalmers. A, young tropho- 
zoite ; B, full-grown trophozoite ; C, early schizont ; 
D, mature schizont ; E, female gametocyte ; F, male 
gametocyte. (After Castellani and Chalmers.) 

Remarks. — Castellani and Chahners (1910) described this 
parasite from dogs in Colombo. Castellani (1924) again 
refers to it, and states that he saw several cases of infection 
in dogs in Colombo. No other observer has seen the organism. 
Wenyon (1926) obtained blood-films from 500 pariah dogs 
from Ceylon, but in none of them could the parasite be found. 

Habitat. — Bloodof the dog, Cams /ami7tam Linn. : Ceylon, 

242. Plasmodium equi Castellani & Chalmers. 

■fPlasmodium equi, Castellani & Chahners, 1913 ; 1919, p. 516. 
Plasmodium equi, Wenyon, 1926, p. 976; Reichenow, 1929, p. 1009 ; 
Coatney & Roudabush, 1936, p. 339. 

Very similar to P. vivax of man. 

Remarhs. — Castellani and Chalmers gave no description, 
but simply stated that the species closely resembled P. canis. 
It has not been recorded since by any other observer. 

Habitat. — Blood of the horse, Equus caballus Linn. : Ceylon. 


243. Plasmodium mackiei de Mello & de Sa. (Fig. 140.) 

■\Plasmodiuni mackiei, de Mello & de Sa, 1916, pp. 736-7, pi. Ixiv, 
fig. C, 1-10. 
Plasmodium mackiei, Wenyon, 1926. p. 1364 ; Coatney & Rouda- 
bush, 1936, p. 339. 

Young trophozoite of a baciUary form. The schizonts show 
a, precocious division of the chromatin, and always form six 
merozoites in the rosette. Gametocytes spherical or ovoid. 

B C 

Fig. 140. — Plasmodium, tnackiei de Mello & de Sa. A, young tropho- 
zoite ; B, schizont ; G, gametocyte. (After de Mello and 
de Sa.) 

Habitat. — Blood of Myotis muricola (Hodgs.) {=Vespertilio 
muricola) : Poktijguese India, Arjuna (Bardez), Santa Cruz. 

244. Plasmodium narayani de Mello & Dias. (Fig. 141.) 

■\Plasmodium narayani, de Mello & Dias, 1936, pp. 212-13, pi. iii. 

Young trophozoite has an annular form, with its cytoplasm 
more or less thin or more or less compact. Sometimes two 
" rings " occur in the same red blood-corpuscle. Parasite 

A B C D E F 

Fig. 141. — Plasm.odium narayani de Mello & Dias. A, flimsy " ring " ; 
B, compact " ring " ; C, two " rings " in the same corpuscle ; 
D, full-grown trophozoite ; E, " rosette " showing schizo- 
gony ; F, female gametocyte. (After de Mello and Dias.) 

not seen in the living condition, but inferred to be strongly 
amoeboid, as many aberrant forms, such as fusiform, crescentic 
or triangular, were met with. Young schizont occupies 
one-sixth to one-fifth of the red cell, and sometimes shows one 
or two granules of almost black pigment. Full-grown schizont 
round, with many granules of pigment, some of them rod- 
shaped, filling the cytoplasm. Chromatin lodged in a clear, 
vacuolated zone, in contrast with the cytoplasm, which is 




more compact. Very rarely " rosettes," with merozoites, 
found in the lung-smears. Female gametocyte oval, with a 
compact oval nucleus situated in the middle and some granules 
and rods of pigment irregularly scattered in the surrounding 
area. Male gametocytes not found. Infected red cell 
sometimes dehsemoglobinized, somewhat enlarged ; but does 
not show dots of any kind. 

Eemarks. — The parasites were very scanty and were found 
in the blood and lung-smears only. 

Habitat. — Blood and smears from the lung of the fish-otter, 
Lutra lutra Linn. : Portfgtjese India, Nagoa. 

245. Plasmodium pteropi Breinl. (Fig. 142.) 

Plasmodium pteropi, Breinl, 1912 ; Johnston, 1913, p. 35, figs. ; 

A. & M. Leger, 1914, pp. 399-401. 
^Plasmodium pteropi, Mackie, 1914, pp. 375-6, pi. xlvii ; Wenyon, 

1926, p. 974, pi. xvi, figs. 8-12, p. 1362. 
Plasmodium pteropi, Knowles, 1928, p. 442 ; Reichenow, 1929, 

p. 1009 ; Coatney & Roudabush, 1936, p. 340. 

Resembles P. vivax of man. Trophozoites ring-shaped, 
showing a circular form when fuU grown. Pigment light 


Fig. 142. — Plasmodium, pteropi Breinl. A, ring-form ; B, trophozoite ; 
G, gametocyte. (After Mackie.) 

brown. Immature schizonts and gametocytes also observed. 
Infected red blood- corpuscles may be sHghtly enlarged. 
Sporogony not known. 

Remarks. — Breinl (1912) described P. pteropi from the blood 
of a flying fox, Pteropus gouldi Peters, in West Austraha. 
Johnston (1913) and A. & M. Leger (1914) described P. pteropi 
as a new species. Mackie (1914) also described a very similar 
form from Pteropus medius Temm. {= Pteropus edwardsii 
Horsfield), and gave it the name P. pteropi without knowing 
that Breinl had already given the name to the West Austrahan 
form. According to Wenyon the two are probably identical. 
The parasite was also seen in the blood of the flying fox in 
Ceylon, and Wenyon (1926) has drawn the figures from these 

Habitat. — Blood of the flying fox, Pteropus medius Temm. 
—P. edwardsii Horsfield) : India ; Ceylon. 



246. Plasmodium ratulse Donovan. 

■^Plasmodium ratufse, Donovan, 1920, pp. 719-20. 
Plasmodium ratufce, Wenyon, 1926, p. 974 ; Knowles, 1928, p. 442 ; 
Reichenow, 1929, p. 1009. 

Very similar to P. vivax of man. Gametocytes predominated, 
and in a few films the female gametocytes exceeded in number 
the male forms. 

Remarks. — The species is similar to, if not identical with, 
P. vassali (Laveran, 1905), the parasite first found by Vassal 
in Sciurus grisemanus in Annam. 

Habitat. — Blood of the giant squirrel, Ratufa indica 
Erxleben : Bombay, Malabar. 

247. Plasmodium tyrio de Mello, Fernandes, Correa, & Lobo. 

(Fig. 143.) 

"fPlasmodium tyrio, de Mello, Fernandes, Correa, & Lobo, 1928, 

pp. 513-16, figs. 1-28. 
Plasmodium tyrio, Coatney & Roudabush, 1936, p. 340. 

Young trophozoites usually circular or oval, very regular ; 
aberrant or fusiform forms very rare. Ring-forms show a 



Fig. 143. — Plasmodium tyrio de MeUo, Fernandes, Correa, & Lobo. 
A, B, ring-forms ; C, D, rosette-forms ; E, F, male gameto- 
cytes ; 6, H, female gametes. (After de Mello, Fernandes, 
Correa, and Lobo.) 

precocious division of the nucleus into two or three granular 
masses united together. Schizonts irregular in form, resemble 
those of P. vivax, and even when fully developed do not 
cover the entire area of the corpuscle. Large black pigment 
granules appear irregularly in the cytoplasm of the parasite. 
Schizogony takes place in the peripheral blood. The number 
of merozoites varies from four to eight. Gametocytes spherical 
or ovoid. Male gametocytes almost devoid of pigment ; 
when the pigment is present, it is in the form of fine dark 
granules dispersed round the nucleus. Female gametocytes 


have a more deeply staining cytoplasm, smaller nucleus, and 
large black granules of pigment. 

Dimensions. — Schizonts l-5-4-5jU, by 1-2|U, ; gametocytes, 
male 3-4-5|u. by 2-2-5 ^m, female 3-4/Lt by 2-2-5/x. 

Habitat. — Peripheral blood, as also smears from lungs 
and hver of the ant-eater, Manes pentadactyla Linn, (popularly 
known in Goa as the " Tyrio ") : Portuguese India, Nova 

3. Family THEILERIID^ du Toit, 1918. 

Parasites of blood of Mammals, which do not form pigment 
(hsemozoin). Schizogony takes place in the endothelial cells 
of the capillaries of the internal organs, and the schizonts 
(Koch's "blue bodies ") produce a number of merozoites. The 
parasites finally invade the red corpuscles, within which they 
occur as round, ovoid, rod-like or irregular forms. Show no 
tendency towards a paired arrangement. The forms in the 
red corpuscles do not reproduce, and are possibly gametocytes. 

The family includes the genus Theileria, the best-known 
species of which, T. parva, causes the East Coast fever of cattle 
in Africa and elsewhere. Species of Theileria have also been 
recorded from sheep and goats in other countries. Du Toit 
(1918) recognizes Eangelia Carini & Maciel (1914) as a distinct 
genus, but Wenyon (1926) has advanced convincing reasons 
for merging the two genera into one. 

Remarks. — There is no teneral agreement as regards the 
position of this family and the following one in the system 
of classification. Wenyon (1926) places the two families 
Theileriidse and Babesiidse in a suborder Pieoplasmidea. The 
suborder includes parasites which inhabit red blood-corpuscles 
of Mammals, but do not form pigment (hsemozoin). In 
films stained with Romanowsky's stain each parasite consists 
of a blue-staining cytoplasm and a red-staining nucleus, 
the latter generally consisting of a granule of chromatin 
with a string of finer granules extending from it. If a vacuole 
is present it becomes difficult to distinguish the form from 
the young ring-form of a malarial parasite. The organisms 
included in the suborder reproduce by division into two or 
four individuals only. 

Reichenow (1929) places the family Theileriidse under 
H^MOSPORiDiA and the family Babesiidse as an appendix 
to the same. In his latest work (1935) he places both the 
families in an appendix after the H^mosporidia, and remarks 
that the life-cycle of the Theileriidse, so far as is known, 
shows points of agreement with the typical H^mosporidia 
but the plan of development of the Babesiidse is quite different 
from the Telosporidia. In this work I have included the 


PiROPLASMiDEA Under H^MOSPOBIDIA on grounds of practical 
convenience, and have divided the order into four famihes 
(see p. 209). 

The two families, Theileriidae and Babesiidse stand in much 
the same relationship to one another as Hsemoproteidse to 

Genus THEILERIA Bettencourt, Franca, & Borges, 1907. 

Piroplasma (part), Stephens & Christophers, 1903 b, pp. 335-6; 

Theiler, 1904, pp. 1-20. 
Theileria, Bettencourt, Franca, & Borges, 1907, pp. 341-9 ; Nuttall, 

Fantham, & Porter, 1909, pp. 325-40 ; Gonder, 1910 a, pp. 143- 

64 ; 1910 b, pp. 49-52 ; 1911 a, pp. 222-31 ; 1911 b, pp. 170-8 ; 

Minchin, 1912, pp. 379-80, 382; Nuttall, 1913, pp. 302-20; 

Hegner & Taliaferro, 1924, p. 305 ; Wenyon, 1926, pp. 992, 1029- 

39 ; Knowles, 1928, pp. 457-9 ; Reichenow, 1929, pp. 1022-7 ; 

Kudo, 1931, p. 289 ; Calkins, 1933, p. 566 ; Reichenow, 1935, 

p. 377. 

With the characters of the family. 

Remarks. — It is not easy to decide whether a particular 
form met with in the blood is a Theileria or a Babesia. In the 
former, schizogony takes place in the endothehal cells of the 
capillaries of the internal organs, and forms produced there 
enter the red corpuscles and are seen in the peripheral blood. 
Unhke Babesia, they do not multiply in the red corpuscles. 
The blood is consequently not infective when inoculated to 
healthy animals imless endothelial cells containing schizonts 
happen to be present. 

248. Theileria parva (Theiler). (Fig. 144.) 

Piroplasma bigemina, a stage of, Koch, 1898. 

Piroplasma kochi, Stephens & Christophers, 1903 6, pp. 335-6, 
fig. 75. 

Piroplasma parvum, Theiler, 1904. 
■\Piroplasma parvum, Lingard, 1907, p. 274, pi. vii, fig. 1, 8. 

Theileria parva, Bettencourt, Franca & Borges, 1907, pp. 341-9, 
pis. xvii, xviii ; Nuttall, Fantham & Porter, 1909, pp. 325-40. 
'\Piroplasma parvum, Gaiger, 1910, p. 66. 

Theileria parva, Gonder, 1910 a, pp. 143-64 ; 1910 b, pp. 49-52 
1911 a, pp. 222-31 ; 1911 b, pp. 170-8 ; Minchin, 1912, p. 382 
Nuttall, 1913, pp. 302-20 ; Castellani& Chahners, 1919, pp. 498-9^ 
fig. 164 ; Hegner & Taliaferro, 1924, p. 305 ; Saceghem, 
1925, pp. 651-60; Wenyon, 1926, pp. 1029-34, fig. 428; 
Sergent, Donatien, Parrot, Lestoquard, & Plantiu-eax, 1926; 
p. 1362 ; 1927 a, pp. 489-506, 3 figs. ; 1927 b, pp. 721-84 
23 figs. ; 1927 c, pp. 161-87, 5 figs. ; Knowles, 1928, p. 457 
figs. 106, 107 ; Reichenow, 1929, pp. 1032-4, figs. 1007-11 
Kudo, 1931, p. 289, fig. 122, i ; du Toit, 1931, pp. 547-8 
Reichenow, 1935, p. 377. 

The Cycle in Cattle. — The sporozoites injected by the tick 
collect in the spleen, lymphatic glands, and other organs, 
penetrate the endothelial cells of the capillaries, grow rapidly, 



and form large multinucleate masses or schizonts (Koch's 
"blue bodies "). They may be discovered by puncture of the 
enlarged glands. In dried films, stained by Romanowsky's 
stain, they will be seen as blue masses of cytoplasm with 
a varying number of red chromatin dots. In films they may 
be found in the endothelial cells or free, but in sections they 
are always intracellular. About fourteen days after infection, 
and simultaneously with the attack of fever, the schizonts 
may be recognized as of two kinds, some containing a smaller 
number of larger nuclei and others containing a larger number 

Fig. 144. 




-Theileria parva (Theiler). A-D, stages in the red blood- 
corpuscles ; E, free schizont ; F, intracellular schizont 
which gives rise to merozoites ; G, free gametocyte-pro- 
ducing body ; H, intracellular gametocyte-producing 
body ; /, yomig gametocytes in a blood-corpuscle ; J, ripe 
microgamete ; K, ripe macrogamete ; L, zygote : M, ooki- 
nete ; N, multiplication cysts in the salivary glands of 
the tick ; 0, free sporozoites. (From Reichenow ; A-D after 
Theiler's preparations ; E-0, after Gonder. ) 

of smaller nuclei. The former break up into a number of 
minute bodies (merozoites), which enter other cells, grow, 
and reproduce by schizogony. The latter give rise to gameto- 
cytes which enter the blood and penetrate the red corpuscles. 
These forms in the blood are very small, ring-, comma-, pear- 
shaped or rod-like ; in the latter the nucleus hes at one end 


Sometimes cross-shaped forms are met with, but these are 
regarded as aggregates of four parasites, and not fission 
stages. The forms in the blood do not multiply by fission, 
and are consequently assumed to be gametocytes. They 
grow into adult gametocytes of two kinds — male gametocytes, 
which are long, slender, " bacillary " forms, and female 
gametocytes, which are plump, rounded or pear-shaped forms. 
The gametocytes can only develop further in the tick, 
Bhipicephaliis . 

Dimensions. — Schizonts 3-10/i in size ; gametocytes, rod- 
like forms 2-5 fjb in. length by l'2fx. 

The Cycle in the Tick. — In the gut of the tick the gametocytes 
leave the red corpuscles and develop into gametes. The 
macrogamete is stationary, whilst the microgamete is motile, 
seeks out the macrogamete, and is said to unite 'wdth it, 
forming the zygote. The latter develops into a worm-like 
ookinete, and no further development is known to take place 
till after the moulting of the tick. After the moulting of the 
tick, large cyst-like structures containing numerous nuclei 
are met with in the salivary glands. Each cyst develops 
a number of sporoblasts, and numerous very small sporozoites 
are ultimately set free, which are introduced into the cattle 
when the tick again sucks the blood. 

Bemarks. — The organism produces a serious disease of cattle, 
known as East Coast fever, in various parts of Africa. It has 
also been met with in Transcaucasia, Macedonia, and India. 
The disease differs from that caused by Babesia higemina in 
that hsemoglobinuria, jaundice, and progressive anaemia are 
absent. Theiler (1904) demonstrated that East Coast fever 
was a distinct disease, and that cattle which had recovered 
from hsemoglobinuric fever due to Babesia were not immune 
to it. He also found that the disease could not be transmitted 
by inoculation of infected blood into a healthy animal. This 
was explained as being due to the fact that the blood-forms 
do not multiply. Experiments by Ed. Sergent and his co- 
workers (1926) have, however, shown that this view is incorrect, 
though, as Wenyon (1926) remarks, the occasional positive 
results obtained by blood inoculations of healthy animals may 
be due to the presence of endothelial cells in the peripheral blood. 

The blood -forms were carefully described by Nuttall, 
Fantham, and Porter (1909) in artificial infections of animals 
in England by means of ticks, Bhipicephalus evertsi, imported 
from South Africa ; and also by Nuttall (1913). The life- 
history was studied by Gonder (1910, 1911 a, 1911 6), whose 
account has been followed above, but certain points lack 

Lingard (1907) and Gaiger (1910) recorded this species 
from India, but no one seems to have seen it since. 

Habitat. — Blood of ox, Bos taurus Linn. (?) : India. 


249. Theileria mutans (Theiler). (Fig. 145.) 

Piropla»ma mutans, Theiler, 1906, pp. 292-300 ; 1909, pp. 115-33. 
■fPiroplasma mutans, Gaiger, 1910, p. 66 ; Baldrey, 1911, p. 569. 
Piroplasma m,utans, Theiler, 1911, pp. 202-3. 
Babesia ^nutans, Minchin, 1912, pp. 380, 382. 
Gonderia tnutans, du Toit, 1918, p. 86, fig. 8. 
Theileria mutans, Castellani & Chalmers, 1919, p. 499 ; Brumpt, 

1923, pp. 16-53, pis. i, ii ; Velu, 1923, pp. 54r-64, 6 figs. ; Doyle, 

1924, pp. 18-27. 

■fTheileria mutans, Edwards, 1925, pp. 48-9 ; 1926, p. 43 ; Cooper, 
1926 a, pp. 96-7 ; 1926 b, pp. 315-16, pi. xviii, fig. 2. 
Babesia mutans, Wenyon, 1926, pp. 1001-2, 1035, fig. 413 ; 

Knowles, 1928, pp. 451, 453, fig. 106. 
Theileria mutans, Reichehow, 1929, p. 1025, fig. 1012 ; Du Toit, 
1931, pp. 551-2. 
■fTheileria mutans. Cooper, 1931, pi. i, figs. 2, 3 ; Achar, 1935, p. 9. 
Theileria tnutans, Reichenow, 1935, p. 377. 

Blood-forms and schizonts very similar to those of T. parva. 
The blood-forms are exceedingly minute, comma-shaped, 
bacilhform or coccal ; dumbbell-shaped, ring or even cross- 
shaped forms occur, the largest not exceeding 1 fj, in diameter. 
Schizonts very similar to Koch's " blue bodies " ; they are 
larger than those of T. parva, and the nuclei tend to be ovoid 


Fig. 145. — Theileria mutans (Theiler). ( x c. 4000.) A-D, different 
forms in the blood -corpuscles. (From Wenyon, after 
Gonder. ) 

and not spherical as in the latter. They occur in small 
numbers in the internal organs as well as in the peripheral 

Transmission occurs by ticks of the genus Bhipicephalus ; 
ticks fed at the nymphal stage transmitted the infection when 
feeding as adults. 

Eemarks. — Compared with T. parva, T. mutans is a benign 
parasite. The infections are mild in character, and the 
parasite does not cause haemoglobinuria, though it may 
cause a certain amount of anaemia. Theiler (1906) showed that 
inoculation of infected blood readily conveys infection to 
healthy animals, and this was regarded as the chief distinction 
between T. parva and T. mutans, so much so that cross-forms 
which occur in both species were differently interpreted, 
in T. parva as being due to aggregation of four parasites, and 
in T. mutans as binary fission, and the latter organism was then 


referred to Babesia. Brumpt (1923) showed that usually the 
infections are mild and the schizonts are not apparent, but in 
intense infections they occiu" in large numbers in the internal 
organs as well as in peripheral blood. This was confirmed 
by Velu (1923), Doyle (1924), and Edwards (1925). Ed. 
Sergent and his co-workers (1924) disagreed -vsdth Brumpt, 
and stated that T. mutans never produces schizonts, but later 
(1928) found a few of them. T. mutans and T. parva cannot be 
differentiated morphologically, and Brumpt (1924) dijfferentiates 
them biologically as follows : — 

Non-pathogenic, transmissible by direct blood inoculation; 
persistence of parasites in the blood for long periods 
(no immunity) T. mutans. 

Highly pathogenic, not transmissible by direct blood inocu- 
lation ; no persistence of parasites after recovery (com- 
plete immunity) T. parva. 

Du Toit (1931) recommends the provisional recognition of 
four species of Theileria in cattle, viz., T. parva, T. dispar, 
T. annulata, and T. mutans, and these show a gradual transition 
from the most virulent T. parva to the avirulent T. mutans. 

Cooper (1926 a) remarked that T. mutans was almost 
universally present in Indian cattle, and showed the occurrence 
of Koch's " blue bodies " in this species. Datta (1938) has 
found that the parasite multiples in the lymphoid tissue 
generally, including that of glands, spleen, etc. 

Habitat. — ^Blood of bulls, Bos indicus Linn. (?), used for 
Rinderpest hyper-immunisation : Punjab ; United Pro- 
vinces, Muktesar ; Mysore, Bangalore. 

250. Theileria sp. 

fTheileria sp., " types A & B," Sen & Srinivasan, 1937, pp. 15-37, 
pi. vii. 

Usually round or shghtly oval forms, having the appearance 
of signet-rings, and uniformly 1-2 /x in diameter. Stained 
with Giemsa's or Leishman's stain, the cytoplasm takes a pale 
blue colour ; the chromatin stains well, two chromatin spots 
being sometimes connected together. Koch's " blue bodies " 

RemarJcs. — Sen and Srinivasan (1937) studied Theileriasis 
of cattle in a large number of Indian hill bulls artificially 
infected with a strain isolated from a fatal case at Muktesar. 
They came to the conclusion that in the present state of our 
knowledge it is impossible to rely on any morphological 
features as an indication of the extent of the pathogenicity 
of the organism. According to them, the diagnosis of the 
species of Theileria would appear to depend on the quantitative 


combination of a number of morphological and clinical features. 
They consider the forms studied by them to be two types of 
a new species and give their differential characters alongside 
those given in du Toit's table of previously known species. 
The examination of blood taken at the first rise of temperature 
frequently reveals the presence of few parasites and Koch's 
" blue bodies " or both, but, as the disease advances, the 
parasites rapidly increase in number and may eventually invade 
50 to 100 per cent, of the red blood-corpuscles, although 
Koch's " blue bodies" may vary in number from " rare " to 
" numerous." 

Sen and Srinivasan also came to the conclusion that the 
infection in imported Friesian bulls is exotic in origin, being 
probably acquired by the animals during the course of their 
voyage to India, and the parasite concerned is Theileria 

Habitat. — Blood of Indian hill buUs, Bos indicus Linn. : 
United Provinces, Muktesar. 

251. Theileria cellii (Castellani & Chalmers). 

■f Babesia cellii, Castellani & Chalmers, 1910. 
Theileria cellii, Castellani & Chalmers, 1919, p. 500. 
Babesia cellii, Wenyon, 1926, p. 1027. 

Bacillary and pear-shaped forms, l3dng side by side in the 
red blood- corpuscles. Development not known. 

Remarks. — This form was originally described as Babesia 
cellii by Castellani and Chalmers, but in the third edition of 
their work they substituted the name Theileria cellii without 
giving any further details. It is by no means clear from the 
original description whether the form should be referred to 
Babesia or to Theileria. 

Habitat.' — Blood of Macacus pileatus (Shaw) : Ceylon. 

252. Theileria hirci Dschunkovsky & Urodschevich. (Fig. 146.) 

Theileria hirci, Dschunkovsky & Urodschevich, 1924, pp. 108-10, 
fig. A, 1-24 ; Wenyon, 1926, pp. 1036-8, fig. 430 ; Reichenow, 
1929, p. 1026 ; Thomson & Hall, 1933, p. 228. 
Theileria ovis, Lestoquard, 1924, pp. 122-8, 15 figs. 
■\Theileria ovis, Edwards, 1926, p. 43. 
Theileria ovis, du Toit, 1931, p. 556. 
■fPiroplasma taylori, Sarwar, 1935, pp. 172-5, pis. v, vi, via, vii. 
Theileria hirci, Reichenow, 1935, p. 378 ; Bhatia, 1936, pp. 151-4. 

Parasites mostly small and variable in shape, some being 
distinctly ring-shaped, others bacillary or nail-like, oval or 
pyriform ; occurring singly, in twos or, rarely, in threes in the 
centre of the red blood-corpuscle. Cross-like forms also occur, 
each member being pyriform. Extracellular forms, similar 
to Koch's " blue bodies," encountered in peripheral blood, 
were round or oval, at times as large or larger than a blood- 


corpuscle, and contained chromatin granules of varying size 
and shape. Internal organs of host contain the schizonts. 

Remarks. — Sarwar (1935) described a form from a goat in 
the Punjab in which the parasites were mostly ovoid or round, 
pear-shaped forms being rare. Single parasites found in 
a corpuscle measured 2^ by 1-5/^. Although single parasites 
were frequently seen, it was not uncommon to find two, 
four, eight or sixteen individuals in one red cell. He also 

E F a 

Fig. 146. — Theileria hirci Dsch. & Urod. A-E, intra- corpuscular forms ; 
F-G, extra-corpuscular bodies showiug multiple division'. 
(After Sarwar.) 

found extracellular forms showing multiple division, but did 
not regard them as Koch's " blue bodies," nor did he find 
them in the internal organs of the host. He named the form 
Piroplasma taylori. I (1936) thought that the form described 
by Sarwar was identical with T. hirci Dsch. & Urod., 1924*. 
Habitat. — Blood of goat, Gapra hircus Linn. : United 
Provinces, Muktesar ; Punjab, Malwale (Sheikhupura). 

4. Family BABESIID^ Poche, 1913. 

Non-pigmented parasites of the red blood- corpuscles of 
Mammals, which multiply in the corpuscle by division into 
two or four. They are of varying size and shape, and usually 
arrange themselves in pairs of pear-shaped individuals. The 
forms in the corpuscles are individuals reproducing asexuaUy, 
but probably some are gametocy-fces. 

Remarks. — Franca (1917, 1918) gave a detailed classification 
of the family and recognized a number of genera. His classi- 
fication was modified by du Toit (1918), who recognized six 
genera, viz., Babesia Starcovici, Nicollia Nuttall, Nuttallia 
Franca, Smithia Franca, Rossiella Nuttall, and Gonderia 
du Toit. Wenyon (1926), Reichenow (1929), and many other 

* Through his courtesy I have now examined his preparations, and 
the extra-corpuscular bodies (fig. 146, F, G) in the peripheral blood 
which linterpreted as Koch's "blue bodies " cannot be regarded as such. 
So his form, ought to be transferred to Babesia and linown as B. taylori 


writers, place all the species in a single genus, Babesia, with, the 
characters of the family. Yakimoff (1931) retains the generic 
name Piroplasma and divides the genus into two subgenera, 
Piroplasma s. str. and Babesiella. Sometimes as many as 
a dozen or more of these parasites occur together in a mammalian 
red corpuscle. They produce no pigment, but destroy the 
corpuscle in which they are contained and set free the haemo- 
globin, which is then excreted by the kidney of the host. 
Hence a characteristic symptom of the diseases produced 
by these parasites, generally termed " piroplasmoses " (or 
" babesioses "), is a great reduction of the corpuscles and 
a red coloration of the urine (hsemoglobinuria or " red- water "). 

Genus BABESIA Starcovici, 1893. 
(JSyn. Piroplasma Patton, 1895.) 

Hiematococcus {non Agardh, 1828), Babes, 1888, p. 692 ; 1890, 

pp. 800, 975 ; 1891, p. 81 ; 1892, p. 359. 
Babesia, Starcovici, 1893, pp. 1—8. 

Pyrosoma (non Peron, 1804), Smith & Kilborne, 1893, p. 67. 
Babesia, Kroguis & Van Hellens, 1894, pp. 353-64; San Felice & 

Loi, 1895, pp. 295-6. 
Apiosoma {non Blanchard, 1885), WandoUeck, 1895, pp. 554-6. 
Piroplasma, Patton, 1895, p. 498 ; Labbe, 1899, p. 124. 
Babesia, Labbe, 1899, p. 125. 
Piroplasma, Laveran, 1901, pp. 385-8 ; Minchin, 1903, pp. 254, 255, 

265, 269, 309. 
Babesia, Minchin, 1903, p. 269. 
Piroplasma, Lingard & Jennings, 1904, pp. 161-5 ; Raymond, 1904, 

p. 272 ; Stephens & Christophers, 1904, pp. 332-7 ; Theiler, 

1904, pp. 401-5; 1906 a, pp. 283-92; 1906 6, pp. 292-300; 

1907, pp. 1-18 ; Dreyer, 1910, pp. 37-45 ; Nuttall & Strickland, 

1910, pp. 524-5 ; 1912, pp. 65-96. 
Babesia, Minchin, 1912, pp. 379-86. 
Piroplasma, du Toit, 1918, pp. 84-104; Castellani & Chalmers, 

1919, pp. 492-8. 
Babesia, Wenyon, 1926, pp. 992-1028; Cooper, 1926 a, pp. 314-15; 

Knowles, 1928, pp. 445-57 ; Reichenow, 1929, pp. 1032-40. 
Piroplasma, Yakimoff, 1931, pp. 372-400. 
Babesia, Kudo, 1931, p. 289 ; Calkins, 1933, p. 566 ; Reichenow, 

1935, p. 378. 

Intra-corpuscular parasites, without pigment. Pyriform 
(lancet-shaped), mostly in pairs, ring-shaped or elhptical. 

The genus includes pa,rasites from dogs, cattle, pigs, sheep, 
horses, rats, mongooses, monkeys, etc., which are transmitted 
by ticks. 

Remarks. — Species belonging to this genus have been de- 
scribed in India or Ceylon from dogs, jackals, horses, cattle, 
goats, mongooses, monkeys, etc. Raymond (1904) was the 
first to draw attention to the presence of indigenous Piroplasma 
in Bo vines in India in 1898. Lingard and Jennings (1904) 
described and figured specimens of Piroplasma from a large 
number of Indian animals and referred them all to a single 


species, Piroplasma tropicus, instead of to several previously 
described species, such as P. bigeminum, P. canis, P. bovis, 
P. ovis, etc. Yakimoff (1931), after reviewing the previous 
classifications of Piroplasma, divides the genus into two 
subgenera, as follows ; — 

(1) Of typical pyriform shape, often in pairs. Length 
of pyriform organisms greater than the radius of the erythro- 
cyte. Situation in the corpuscle central. Pyriform indi- 
viduals of a pair form an acute angle. Number of chromatin 
masses in budding pear-shaped forms not less than two. 
Pear-shaped forms more numerous than the round forms. 
Trypan blue effective. Species included are P. bigeminum, 
P. caballi, P. canis, P. motasi \_=^P. avis']. Subgenus Piro- 
plasma s. str. 

(2) Form pear-shaped (lancet-shaped), annular or elhptical. 
Length of the pear-shaped forms smaller than or equal to the 
radius of the erythrocyte. Situation in corpuscle peripheral 
or central. Pyriform individuals of a pair form an obtuse 
angle. In the pear-shaped forms there is a single chromatin 
mass. Round forms predominate over the pear-shaped ones. 
Trypan blue may or may not be effective. Subgenus 
Babesiella Mesnil, 1919. 

The subgenus Babesiella is further subdivided into species 
groups as follows : — 

Group 1. Babesiella s. str. Form smaller than the radius 
of the erythrocjrte ; situation peripheral ; trypan blue 
effective B. bovis. 

Group 2. Frangaiella. Form smaller than or equal to 
the radius ; situation central ; trjrpan blue not effective. 

(a) Form equal to radius . . F. colchica, F. major. 
(6) Form less than the radius. F. argentina, F. berbera, 
F. caucasica, F. occidentalis. 

This scheme is convenient for diagnostic purposes, but 
obviously the names Babesiella and Frangaiella cannot be used 
in a generic sense. 

253. Babesia bigemina (Smith & Kilborne). (Fig. 147.) 

Pyrosoma bigeminum. Smith & Kilborne, 1893 a, p. 67 ; 1893 b, 

p. 511, fig. 1 ; Starcovici, 1893, pp. 1-8. 
Piroplasma bigeminum, Labbe, 1899, p. 194, fig. 195 ; Minchin, 

1903, pp. 242, 262, 269, 338, 350. 
■^Piroplasma bigeminum, Lingard, 1903. 

"^Piroplasma sp. (probably bigeminum), Donovan, 1903, p. 1401. 
■\Piroplasma bovis, Stephens & Christophers, 1904, pp. 333-4, 

pi. iii. 
■\Piroplasma bigeminum, Hohnes, 1904, pp. 317-26, 1 pi. ; Lingard 
& Jennings, 1904, pp. 161-5, pi. i, figs. 4, 5 ; 1907, pi. vii, fig. G, 9. 
Piroplasma bovis, Nuttall & Graham-Smith, 1908, pp. 138-9, 
diagrana ii. 



■fPiroplasma bigeminum, Gaiger, 1910, p. 66 ; Baldrey, 1910, 

pp. 569-79. 
■\Piroplasma {Babesia) bigeminum, Edwards, 1925, pp. 48-9. 
Piroplasm.a bigeminum,, Minchin, 1912, p. 379, fig. 160 ; pp. 384—5, 

fig. 162 ; Castellani & Chalmers, 1919, p. 497 ; Velu, 1922, 

pp. 133-45, fig. 21 ; Hegner & Taliaferro, 1924, p. 304. 
■\Piroplasma bigem,inum,. Cooper, 1926 a, p. 96 ; 1926 b, pp. 314- 

15, pi. xviii, fig. 1. 
Babesia bigemina, Wenyon, 1926, pp. 993-8, figs. 408, 410 A-J, 

pi. xviii, figs. 6-10; Knowles, 1928, p. 451, figs. 105, 106; 

Reichenow; 1929, pp. 1034-5, fig. 1019 ; Dennis, 1930, pp. 179- 

92, 2 pis. ; Kudo, 1931, p. 289, fig. 122, g. 
-fEabesia bigem,ina. Cooper, 1931, pi. i, fig. 1 ; Ware, 1932, p. 31 ; 

Ray, 1938, p. 265. 

Largest of the known Piroplasms of cattle. Round, oval 
or irregular, or pyriform and occurring in pairs, individuals 
of the pair lying close together : the pear-shaped forms 
extend across the diameter of the red corpuscle, which in cattle 
measures from 5 to 6/i,. Occasionally four pear-shaped forms 
are arranged in a fan-hke manner. MultipHcation takes place 

Fig. 147. — Babesia bigemina (Smith & Kilborne). ( X c. 3000.) 
A, pear-shaped form ; B-F, multiplication by budding ; 
G, two pear-shaped forms in a corpuscle ; H, four pear- 
shaped forms in a corpuscle. (From Wenyon, after Nut tall 
and Graham-Smith.) 

by a characteristic budding process, the buds remaining 
attached by their pointed ends. Transmission by ticks. 

Dimensions. — Round forms 2-3 />t, in diameter ; pyriform 
examples 2-4 ju, in length by 1-5-2/x in width. 

Remarks. — It was in the case of B. bigemina, the cause of 
red- water fever in cattle in Texas, that Smith and Kilborne 
(1893) demonstrated the possibility of transmission of proto- 
zoal parasites by Arthropod hosts. Not only did they discover 
that infection is transmitted by the tick, but that the eggs 
of an infected tick are also infected, so that the second generation 
of ticks hatched from such eggs may also be infective. This 
was the first time that the role of Arthropods in the trans^ 
mission of protozoal disease was demonstrated — several 


years before Ross's discovery of the transmission oi Plasmodium 
prsecox by Culex mosquitoes. Not much, however, is known 
of the actual development in the tick. Koch (1906 a) observed 
that in the stomach of the tick the pear-shaped forms escaped 
from the blood-corpuscles, became amoeboid, threw out long 
spiky pseudopodia, associated in pairs, and gave rise to elongate 
bodies with two nuclei and spiky pseudopodia at both 
extremities. Fusion of the nuclei then took place, along 
with the withdrawal of the pseudopodia. Christophers 
(1907), who studied the development of B. canis in the tick, 
doubted if the amoeboid stages were essential in the develop- 
ment, and described motile club-shaped forms, which he 
referred to as " zygotes " or " ookinetes." These give rise 
to a globular stage, which divides by multiple fission into 
sporoblasts and sporozoites. The sporozoites collect in vast 
numbers in the salivary glands of the tick, and thence pass into 
the Vertebrate when the tick feeds. Observations on the 
development in cultures were pubhshed by Kleine (1906) and 
by Nuttall and Graham-Smith (1908), and certain stages in 
the tick were also described by Dschunkovski and Luhs (1909). 

The development of the parasite in the Vertebrate host 
consists of multiplication by binary or quadruple fission 
within the corpuscle. After destroying the corpuscles in 
which they are lodged, the parasites become free in the blood 
and penetrate other red corpuscles. The stages in the blood 
were studied in great detail by Nuttall and Graham- Smith 
(1906, 1907, 1908) and by Christophers (1907) in B. canis, 
and a summary of their observations will be found under that 

The organism remains in the blood of animals, that have 
recovered from acute symptoms, for many years after its 
apparent disappearance, as tested by microscopic examina- 
tions, and inoculation of blood can produce infection in 
a healthy animal. 

Cooper (1926 a) is of the opinion that infection with this 
parasite is so widespread in India that probably in most 
localities cattle become infected as young calves, at which 
age they possess a high degree of resistance and recover 
readily, and are subsequently immune, but continue to act 
as carriers. Acute disease is liable to occur when adult 
cattle are imported from countries where the species does not 
exist, or when the immunity is broken down through the 
effect of intercurrent disease conditions. 

Dennis (1930) described the structure of the nucleus of 
B. bigemina and even noted the presence of a blepharoplast. 

Ray (1938) found the nuclear chromatin to be confined to 
the apical portion of the parasite, with a circular row of fine 



chromatin beads connected with the main mass of chromatin. 
He did not find any evidence of a blepharoplast. 

Habitat. — Blood of ox, Bos indicus Linn. (?) : Punjab, 
Lahore ; United Provinces, Muktesar ; Madras. 

254. Babesia bovis (Babes). (Fig. 148.) 

Hmmatococcus bovis. Babes, 1888, p. 692 ; 1890, pp. 800, 975. 
Babesia bovis, Starcovisi, 1893, pp. 1-8 ; Labbe, 1899, p. 125. 
Piroplasma divergens, MacFadyean & Stockman, 1911, p. 340 ; 

Nutall, 1913, p. 305, fig. 2. 
Piroplasma bigeminum {Babesia bovis), Minchin, 1912, p. 379. 
Microbabesia divergens, Sohns, 1918. 
Babesiella bovis, Mesnil, 1919. 

Babesia bovis, Wenyon, 1926, pp. 998-1000, figs. 411, 412; 
Reichenow, 1929, pp. 1035-6, fig. 1021 ; 1935, p. 378. 
^Babesia bovis, Idnani, 1938, p. 265 ; 1938, p. 42. 

Smaller than B. bigemiTia. Amoeboid forms chiefly rounded, 
often ovoid, pear-shaped, or rod-shaped forms also occur. 

Fig. 148. — Babesia bovis (Babes). (After Nuttall.) 

In films stained with Romanowsky's stain the nucleus is 
seen as a red dot, with a string of fine granules extending from 
it. Two pear-shaped individuals often lie with their pointed 
ends together and forming an obtuse angle, or even in a line. 
The forms are smaller than the radius of the corpuscle, usually 
He near the margin of the corpuscle, and trypan blue is usually 

Dimensions. — Round forms 1-1-5 jn in diameter ; pyriform 
individuals 1*5-2 ju, in length. 

Remarks. — The organisms were marginal in position and the 
pointed ends of a pair of individuals included an obtuse 
angle, or the individuals were in a straight line. The host 
had shown acute piroplasmosis associated with hsemoglobi- 
nuria, and trypan blue had been administered, but without 
producing any effect. 

Habitat. — Blood of an Indian buffalo, Bos bubalus Linn. : 
Madras, Belgaum. 



255. Babesia eaballi (Nuttall & Strickland). (Fig. 149.) 

Piroplasma sp., Marzinowsky & Bielitzer, 1909, pp. 28-31, pis. v— 

Piroplasma caballi, Nuttall & Strickland, 1910, pp. 524r-5 ; 1912, 

pp. 65-96, pi. iii, 8 diagrams and 5 charts ; Dschiinkovski 

& Luhs, 1913, pp. 289-302, pis. xiv, xv ; Darling, 1913, pp. 197- 

203, pi. iii ; Carpano, 1913 b, pp. 13-41, 3 pis. 
■\Piroplasma caballi, Valladares, 1914, pp. 88-94. 
Piroplasma caballi, Yakimoff, Schokhor, & Koselkine, 1917, 

pp. 302-11 ; Vein, 1922, pp. 197-212, fig. 27. 
Babesia caballi, Wenyon, 1926, p. 1008, pi. xviii, 21-25, & fig. 415 ; 

Knowles, 1928, p. 453, fig. 105, 22-30; Reichenow, 1929, 

pp. 1036-8. 

Parasites relatively large, and resemble B. bigemina found 
in cattle. Multiplication takes place by budding, and pear- 
shaped forms occur in pairs in the red blood- corpuscles. 

-Babesia caballi (Nuttall & Strickland). A, pear-shaped 
form ; B-D, stages in the process of budding ; E, a pair 
of pear-shaped individuals. (After Nuttall and Strick- 

Remarks. — This form, like B. equi (Laveran, 1901), causes 
hsemoglobinuric fever in horses. Nuttall and Strickland 
(1910) showed that two distinct species occur in horses. 
They placed the small form, previously known as Piroplasma 
equi Laveran, in a new genus under the name Nuttallia equi, 
and described the larger form as Piroplasma caballi. Later 
(1912) they described the method of multipHcation of the latter 
in the blood. 

Marzinowsky and BieUtzer (1909) showed that the tick, 
Dermacentor reticulatus Koch, was the vector in South Russia, 
and Carpano (1913 b) thought that Margaropus annulatus 
(Say) was the vector in Italy. 

Habitat. — Blood of the horse, Equus caballus Linn. : 
Madras, Madras. 

256. Babesia canis (Plana & GalH-Valerio). (Figs. 150, 151.) 

Pyrosoma bigeminum var. canis, Piana & Galli-Valerio, 1895, 

p. 345. 
Piroplasma canis, Labbe, 1899, p. 124 ; Lounsbtuy, 1901, p. 714 ; 

Minchin, 1903, pp. 242, 270, 337, 350 ; Lounsbury, 1904, p. 113 ; 

Nuttall, 1904, pp. 219-57, 7 figs. & 7 charts. 




■\Piroplasina canis, Daggetty, 1904, pp. 67-8 ; Lingard & Jennings, 
1904, pp. 161-5, pi. i, fig. 10 ; Stephens & Christophers, 1904, 
pp. 334-5, fig. 74. 
Piroplasma canis, Nuttall & Graham- Smith, 1905, pp. 237-49 ; 
1906, pp. 586-651, pis. xi-xiii and 23 text-figs. ; 1907, pp. 232- 
72, pis. i-iii. 
^Piroplasma canis, Webb, 1906, p. 1; Christophers, 1907 o, 

pp. 76-8 ; 1907 6, pp. 1-83, 3 pis. 
Piroplasma canis, Nuttall & Graham-Smith, 1909, pp. 211-14, 

pi. ix. 
■^Piroplasma canis, Baldrey, 1910, pp. 569-79 ; Gaiger, 1910, p. 66 ; 
Brandford, 1912, pp. 643-6. 
Babesia canis, Schuberg & Reichenow, 1912, pp. 415-34, pi. ii. 
Piroplasma canis, Minchin, 1912, pp. 382-3, 384-5, 387, figs. 161, 

"^Piroplasma canis, deMello, de Sa, de Sousa, Dias, & Noronha, 1917, 

p. 16. 
Piroplasma canis, Castellani& Chalmers, 1919, pp. 493-6, fig. 163 ; 
Velu, 1922, pp. 224-40, figs. 29, 30 ; Hegner & Taliaferro. 1924, 
pp. 303-4, fig. 114. 
■^Piroplasma canis, Rau, 1926, pp. 243—4. 
Babesia canis, Wenyon, 1926, pp. 1012-20, pi. xviii ; figs. 410, 417, 
418, 419, 421; KJaowles, 1928, pp. 446-50, figs. 103, 104; 
Reichenow, 1929, pp. 1032-4, figs. 1016, 1018. 
■\Piroplasma [Babesia) canis, Stirluig, 1929, pp. 647-53. 
Babesia canis, BeKtzer & Markoff, 1930, pp. 498-601 ; Regendanz, 
1932, pp. 745-8 ; Regendanz & Reichenow, 1933, pp. 50-71, 
2 pis., 1 fig. ; Reichenow, 1935, p. 378, fig. 31. 

The Cycle in the Dog. — Parasites typically pear-shaped, 
rounded and bulbous at one end and pointed at the other, with 

Fig. 150. — Babesia canis, development in the dog ( x c. 3000). A, pear- 
shaped form ; B, vacuolated form ; C, rounded form ; 
D-G, stages in the process of budding ; H-J, ultimate 
formation of pear-shaped individuals. (After Nuttall and 
Graham-Smith. ) 

a vacuole in the cytoplasm ; 4-5-5;u, in length. In films 
stained with Romanowsky's stain the nucleus is seen as a 
deeply stained granule , situated near the pointed end, with 
a string of fine granules extending from it. This tj^ical 



form passes through a series of stages before division takes 
place. The organism becomes romided, the vacuole first 
enlarges, then disappears, and the string of chromatin granules 
is withdrawn into the larger granule (fig. 150, A-G). It now 
passes through an amoeboid phase, ajid division of the organism 
is initiated by the separation from the larger granule of 
chromatin of a smaller one, which remains connected with the 
larger by a string of granules. The smaller granule divides into 

Fig. 151. — Babesia canis, development in the tick (x c. 1500). 
A, globular forms in the stomach of the tick ; B, globular 
form showing split ; C, club-shaped forms ; D, " zygote," 
which passes through the stomach-wall ; E-H, globular 
body in ovum or tissue-cells, which shows multiple fission ; 
I, sporoblasts ; J, formation of sporozoites ; K, sporozoites 
in the salivary glands. (From Nuttall, after Christophers.) 

two, and the cytoplasm forms two buds, each containing 
one of these granules ; thus finally a trilobed form results, 
in which the three chromatin granules are connected with 
a Y-shaped or V-shaped chromatin strand (fig. 150, D-G). 


The larger chromatin granule also now divides, followed by 
the division of the cytoplasm, and two pear-shaped merozoites 
result (fig. 150, H-J). These merozoites may now leave the 
corpuscle to invade other corpuscles or may further divide 
within the same corpuscle so as to produce multiple-cell 
infections. Schizogony is thus by budding, and is not com- 
parable with that of the malarial parasites. Sometimes 
there may be as many as sixteen of these organisms in a single 
blood- corpuscle . 

The Cycle in the Tick. — When ingested into the stomach 
of the tick the parasites leave the blood- corpuscle, increase 
in size, and become globular bodies, 4-5 /a in diameter. A spHt 
appears in the globular body, a portion swings round, and the 
globular body is changed into a club-shaped body. The club- 
shaped bodies are motile and Gregarine-hke. Whether any 
sexual process is involved is not known. The club-shaped 
bodies, after passing through the wall of the gut, enter the 
ova and later are found in the tissue -cells of the embryo 
developed from the egg. There they again become globular, 
and increase in size up to a diameter of 25 /x. This globular 
stage, termed " zygote " by Christophers, according to 
Minchin probably corresponds to the oocyst of the Plasmodiidae. 
The globular body divides up by multiple fission into a number 
of " sporoblasts," which do not remain aggregated together, 
but scatter themselves through the tissues of the tick, larva, 
nymph, or adult as the case may be. The nucleus of each 
sporoblast divides into a large number, and then the sporoblast 
segments into an equal number of sporozoites, which are 
small bodies with a single nucleus similar in appearance to 
the forms in the blood. The sporozoites collect in the sahvary 
glands of the tick and pass into the blood of the dog when the 
tick next feeds on it. 

Remarks. — The parasite was first discovered as the cause of 
malignant jaundice of dogs in Italy, and has since been shown 
to have a wide distribution. It has long been known to yield 
to treatment with the aniHne dye, trypan blue. Lingard and 
Jennmgs (1904), James (1905), Webb (1906), and Christophers 
(1907 a, 1907 h) were the earliest to study it in India. Christo- 
phers showed that in India JRhipicephalus sanguineus (LatreUle) 
was the transmitting tick, and described the cycle in the tick 
as summarized above. Christophers also proved that eggs 
laid by an adult which had fed on infected dogs gave rise 
to larvae which were not infective, but that the nymphs, 
and probably the resulting adults, were infective. James 
took specimens of R. sanguineus to England and succeeded in 
infecting English dogs with them. Baldrey (1911) gave 
a useful summary of the observations on piroplasmosis in 
India up to that date. 


Lounsbury (1901, 1904 a) was the first to demonstrate 
experimentaUy that infection is transmitted from dog to dog 
by the tick HaemapJiysalis leachi (Audouin) in South Africa. 
Nuttall (1904) and NuttaU and Graham-Smith (1905, 1906) 
described the structm-e and hfe-history of the parasite in the 
dog. Breinl and Hindle (1908) described a biflagellate stage 
of the parasite, but Wenyon (1926) and Knowles (1928) are 
of the opinion that this must have been a species of Bodo 
(llastigophora) from some extraneous source, and was not 
part of the Hfe-cycle of B. canis. Schuberg and Reichenow 
(1912) concluded from their observations that the amoeboid 
forms are extra-corpuscular, become rounded, and produce 
buds, and these buds pass into the red corpuscles, producing 
there the intra-corpuscular pairs of pear-shaped forms. They 
also studied the details of nuclear di^'ision in wet fixed 

Nawrotzky (1912) infected dogs by introducing infected 
blood into the stomach by means of a stomach tube. Laveran 
and Nattan-Larrier (1913) adduced evidence to show that the 
viruses of France and North Africa were different, dogs which 
had recovered from infections with French virus, and were 
immiuie, could be infected with the North African one. 

Several workers have attempted to infect animals other 
than dogs, but without success. Dunsbury (1903) failed to 
infect a jackal. Nuttall and Graham-Smith (1909 a) failed 
to infect foxes. Contrary to these negative results, Rau 
(1926) has succeeded in experimentally infecting a jackal with 
B. canis from a dog. He also found the parasite in a blood- 
smear from a wild jackal. 

Regendanz and Reichenow (1933) have restudied the develop- 
ment of Babesia canis in Dermacentor reticulatus. They 
find that most of the parasites when sucked in with the dog's 
blood by the tick die, but some produce small vermiform 
bodies that infect the cells of the tick's midgut and multiply 
there asexually. The daughter cells enter the body-cavity 
and proceed to the eggs, where, after a few divisions, they 
decrease in size and remain dormant. Later, in the nymph 
or the adult developing from the egg, they enter the salivary 
glands, multiply and produce vermiform cells that escape 
into the saHvary ducts ; thence they are passed into the dog. 
In the dog's blood they reproduce asexually. No sexual 
stages were found by them at any stage of the hfe-history, 
and they beheve that the sexual stages previously reported 
by others are misinterpretations of the observations. They 
further claim that piroplasms have no close relationship 
with either Sporozoa or Flagellata, but have their nearest 
affinities with Sarcodina. 


Habitat. — Blood of the dog, Canis familiar is Linn. : Madras, 
Madras ; Punjab ; Porttjguese India, Nova Goa ; Central 
Provinces ; the jackal, Canis aureus Linn. : Madras, Madras ; 
and the blood of the transmitting tick, Bhipicephalus san- 
guineus Koch. 

257. Babesia equi (Laveran). (Fig. 152.) . 

Piroplasma equi, Laveran, 1901, pp. 385-8, text-figs. ; Minchin, 

1903, pp. 242, 270, 350; Stephens & Christophers, 1904, 

p. 337. 
"fPiroplasma sp., Lingard & Jennings, 1904, pp. 161-5. pi. i, figs. 2, 

Plasmodium equi, Axe, 1906, pp. 222-5 ; Jolifle, 1907, pp. 51-66. 
Piroplasm,a equi, Gaiger, 1910, p. 66. 
Nuttallia equi. Franca, 1910, p. 14 ; Nuttall & Strickland, 1910, 

pp. 524-5 ; Minchin, 1912, p. 380 ; Dschunkvoski & Luhs, 1913, 

pp. 289-302, pis. xiv, xv. 
Piroplasma equi, Carpano, 1913 6, p. 845; 1914 a, pp. 13-41, 

3 pis. ; 1914 b, pp. 42-53, 1 pi. 
"fPiroplasm-a equi, Valladares, 1914, pp. 88-94 ; Williamg, 1914, 

pp. 1-6. 
Nuttallia equi, Schein, 1917, pp. 871-2 ; YakimofE, Schokhor, & 

Koselkine, 1917, pp. 302-11; Du Toit, 1919, pp. 84-104; 

Castellani & Chalmers, 1919, p. 500. 
Piroplasma equi, Velu, 1922, pp. 197-212, fig. 28. 
Babesia equi, Wenyon, 1926, pp. 1009-10, fig. 416, pi. xviii, 

figs. 26-30 ; Knowles, 1928, p. 453, fig. 106, 26-30 ; Reichenow, 

1929, pp. 1036-7, fig. 1022. 

Very small, nearly rounded and actively amoeboid, ovoid 
or pear-shaped, not exceeding 2/x in diameter. Division 

Fig. 152. — Babesia equi (Laveran). (After Wenyon.) 

gives rise to four individuals, which are arranged in a cross- 
hke manner. The four daughter individuals eventually sepa- 
rate, escape from the red corpuscle, and infect other corpuscles. 
Transmission by ticks. 

Remarks. — Round or ring-shaped, oval or rod-shaped piro- 
plasms showing division rosettes of four, arranged in the 
form of a cross, were placed by Franca (1910) in a separate 
genus called Nuttallia : but Wenyon (1926) thinks that 
differences in size and in the number of daughter individuals 
produced are not sufficient grounds for recognition of new 


Nuttall and Strickland (1910) have described the growth 
and division stages in this species. The species produces 
a disease Hke that caused by B. caballi, but is much more 
widely distributed. Theiler (1905 a) showed that Ehipi- 
cephalus evertsi Neumann is the vector in South Africa, and 
according to Carpano (1913 6), Ehipicephalus bursa Canestrini 
& Fanzega is the vector in Italy. 

Young animals are not so seriously affected as older ones. 
Immunity is acquired. The blood remains infective for many 
years after a clinical recovery. 

Habitat. — Blood of the horse, Equus caballus Linn. : Punjab ; 
United Provinces, Muktesar ; Rajputana ; Madras. 

258. Babesia felis Davis. (Fig. 153.) 

"fPyroplasma sp., Lingard & Jennings, 1904, p. 161. 
Babesia felis, Davis, 1929, pp. 304, 523-34. 
■f Babesia felis, Mangrulkar, 1937 a, p. 15 ; 1937 b, pp. 243-6, pi. xvi. 

Small, rounded, non-pigmented, intra-corpuscular parasite 
multiplying by division into four in a cross-like arrangement. 

Fig. 153. — Babesia felis Davis. Various forms seen in Leishman- 
stained films. Light blue cytoplasm is indicated by- 
stippling and the dark red chromatin by black. Outlines 
of the corpuscles are represented schematically. (After 

Schizonts do not occur in the internal organs. Trypan blue 
not effective. 

Dimensions. — 1-2-25/x in diameter, the majority being 
about 1-25/x.. 

Remarks. — Lingard and Jennings (1904) recorded piro- 
plasmosis in wild and tame cats, but did not describe the 
form found. Davis (1929) described Babesia felis from 
the Sudanese wild cat, Felis ocreata, and found that the 
parasite is readily transmissible to the domestic cat by means 
of blood inoculation, and that the progress of infection in 
inoculated cats follows a constant but benign course. Splenec- 
tomy prior to or after inoculation results in an intense 
infection characterized by anaemia and hsemoglobinuria. 
Mangrulkar (1937) has recently found the form in Indian cats, 
but has not come across any division into four or cross-forms. 

Habitat. — Blood of cat, Felis domesticus Linn. : United 
Provinces, Bareilly, Muktesar. 



259. Babesia gibsoni (Patton). (Fig. 154.) 

■fPiroplasma tropicus (part), Lingard & Jennings, 1904, pp. 161-5. 
\Piroplasma gibsoni, Patton, 1910, pp. 274-81, fig. 1. 
"fPiroplasma tropicus (part), Baldrey, 1910, pp. 572—7. 
■fPiroplasma gibsoni, Patton, 1911, pp. 615-21, 1 fig. ; Symons & 
Patton, 1912, pp. 361-70. 
Achromaticus gibsoni. Franca, 1917, pp. 221—9. 
Piroplasma gibsoni, Castellani & Chalmers, 1919, p. 497. 
Babesiella gibsoni, Velu, 1922, p. 131. 
Babesia gibsoni, Wenyon, 1926, pp. 1020-1, fig. 422. 
Piroplasma gibsoni, Symons, 1926, pp. 293-315. 
fPiroplasma gibsoni, Rau, 1927, pp. 785-800, pis. xxxi, xxxii, 

4 text-figs'. ; Stirling, 1929, pp. 647-53. 
f Babesia gibsoni, Swaminath & Shortt, 1937, pp. 499-503 ; Ray & 
Idnani, 1938, p. 265. 

Smaller than B. canis ; young trophozoites rounded, not 
pear-shaped, and appear as small ring-like forms occupying 
about one-eighth of the infected corpuscle. Parasites (in 
films stained with Leishman's or Giemsa's stain) appear 
as small rings, somewhat resembling the rings of the malignant 
tertian malarial parasites but very much smaller and staining 

C J> 

Fig. 154. — Babesia gibsoni (Patton). 
(From Wenyon, after Patton.) 

less deeply. They also differ from B. canis in not showing 
reticular structure of protoplasm or vacuoles. The protoplasm 
takes a very faint blue stain, the ring being sometimes made 
up of a blue-staining periphery and a clear centre. Generally 
there are two masses of compact chromatin, staining red, 
one larger and the other smaller, which is situated either 
near the larger or on the periphery of the ring. Larger 
ovoid or elongate forms are also met with, though in smaller 
numbers. Parasite usually excentrically situated in the blood- 
corpuscle. Usually only one, but sometimes up to five, may 
be found in a single corpuscle. The infected host-cell generally 
not altered in shape or size. Sometimes the parasite escapes 
from the corpuscle, becomes disc-shaped, and after a short 
while enters another corpuscle. In spleen -puncture smears 
similar forms are met with, and also very large rings with their 
chromatin divided into two, three or even four parts. Schizo- 
gony takes place in the spleen. The parasite, having penetrated 
a red cell in the spleen, grows and divides into two, four, etc. 
The daughter parasites are somewhat pear-shaped, and, when 


present in large numbers in a cell, the shape varies from 
a long oval to pyriform or circular. The merozoites are set 
free by the rupture of the corpuscle and infect other corpuscles. 

Remarks. — Patton (1910) believed, that the hounds got the 
infection from the jackal. He demonstrated the same 
parasite in the blood of the jackal, and showed that the 
infection could be transferred to dogs my means of inoculation 
of blood from the infected animal. Dogs which had recovered 
from B. canis were inoculable with B. gibsoni. He suspected 
Bhipicephalus sp., a species related to Rhipicephalus simus 
(Koch) which occurs on the jackal, as the transmitting agent. 
Baldrey (1911) thought that the outbreak described by 
Patton as due to this species was of the same nature as that 
previously described by Pease and Gunn (1908), and that the 
form is not morphologically distinct from Babesia tropicus 
(Lingard & Jennings, 1904). Lingard and Jennings studied 
piroplasmosis in different animals, but probably confused 
a number of species, and B. tropicus is not now recognized 
as a distinct species. 

Rau (1927) observed that smears from engorged larvae of 
the tick, Hssmaphysalis bispinosa Neumann, showed some 
parasites with fibrillar prolongations and a few free large 
rings. Transmission experiments with H. bispinosa were 
inconclusive. Cultivation of the parasite in vitro was also 
not successful. The blood from an infected jackal when 
injected into a dog gives the disease to it and vice versa. 
In the dog, however, the disease takes a more acute form than 
in the jackal. 

According to Stirling (1929) the infection caused by this 
much smaller piroplasm differs clinically from that caused 
by P. canis in being usually more prolonged in its effects. 
The symptoms are generally those of a progressive anaemia 
with frequent tendency to relapses at prolonged intervals. 
It does not respond to treatment with trypan blue, but good 
results have been obtained by repeated treatment with certain 
arsenical preparations, notably tryparsamide. 

Swaminath and Shortt (1937) have shown that the jackal- 
tick, Hsemaphysalis bispinosa Neumann, is a vector of 5. gibsoni, 
and that all stages of the tick can transmit. Hereditary 
transmission through the egg also occurs. 

Ray and Idnani (1938) have recently made a detailed study 
of B. gibsoni in the Vertebrate host. They found two types 
of parasites, viz., ring forms and thin elongate forms, in 
smears from the peripheral blood as well as internal organs. 
The ring forms were found to multiply by repeated binary 
fission until 12 to 16 merozoites were formed ; while the 
thin elongate forms were observed to multiply in a manner 
which suggested a process of schizogony and gave rise to 

316 - SPOROZOA. 

more than 30 merozoites in a corpuscle. Sometimes corpuscles 
infected with both forms were seen in smears from peripheral 
blood, and they regard this as suggestive of sexual dimorphism. 
Habitat. — Blood of the jackal, Ganis aureus JAnn. : Ceylon (?) ; 
Madras, Kurnool ; blood of the dog, Canis familiaris Linn. : 
Ceylon (?) ; Madras ; Central Province. Body of 
Hsema'physalis bispinosa Neumann : Madras, Madras. 

260. Babesia motasi Wenyon. (Fig. 155.) 

Babesia bigemina, Motas, 1903 a, pp. 1522-4 ; 1903 b, pp. 501-4. 
Piroplasma ovis, Dschunkovski & Luhs, 1909, pp. 149-51 ; 

Lestoquard, 1925, pp. 140-5. 
Babesia motasi, Wenyon, 1926, pp. 1005-6, fig. 414, 9-12. 
■[Babesia motasi, Aehar & Srikantiah, 1934, pp. 1-3, 2 pis. & 2 charts. 
Babesia motasi, Sarwar, 1935, pp. 171-2 ; Thomson & Hall, 1933, 

pp. 218-220, figs. 1-6. 

Parasites are most frequently pear-shaped, though ovoid, 
round or irregularly shaped forms are also met with. They 
occur singly or in pairs, and the pairs always meet at an acute 

155. — Babesia motasi Wenyon. ( X 3000.) 
(From Wenyon, after Lestoquard.) 

angle at their narrower ends. Chromatin often appears as 

Dimensions. — 2-5-4/x in length by 1-2-3 /a in breadth. 

Remarks. — Motas (1903) found this parasite in sheep and 
considered it to be identical morphologically with B. bigemina 
of cattle. He also described the transmission of the infection 
through the agency of the tick, Bhipicephalus bursa Canestrini 
& Fanzega. The disease caused by this parasite also resembles 
red-water fever of cattle, and is said to occur in acute or 
chronic form. Lestoquard (1925) observed this parasite in 
the blood of goats in Algeria, and showed that the parasites 
were readily inoculable from one goat to another. He referred 
to it as Piroplasma ovis, but as the name Babesia ovis is 
already preoccupied for a piroplasm of sheep of intermediate 
size, Wenyon (1926) re-named it B. motasi. 

Achar and Srikantiah (1934) have recorded the finding of 
this species in sheep. They have conducted a few transmission 
experiments, and report that the parasite was not transmissible 
to goats. 

Habitat. — Blood of sheep, Ovis sp. : Mysore State. 


•261. Babesia ninense (Yakimojff). (Fig. 156.) 

Piroplasma ninense, Yakimoff, 1909, pp. 472—7, 1 pi. 
^Nuttallia ninense, Sinton, 1922, pp. 359—63, pis. xxvi, xxvii. 
Babesia ninense, Wenyon, 1926, p. 1358. 
Nuttallia ninense, Reichenow, 1929, p. 1039. 

Parasites occur in the red blood-cells, usually one in a cell, 
occasionally more. Stained with Giemsa's stain the cytoplasm 
is blue and the nucleus a deep carmine-red. The cytoplasm 
is condensed and deeply stained at the periphery, with usually 
a hghter area in the neighbourhood of the nucleus. Cyi;o- 
plasm is voluminous as compared with the nucleus, which is 
a soHd mass of chromatin, usually round, but elongated in 
dividing forms, and is usually peripheral, rarely central. The 
intracellular parasites may be ring-shaped, oval or elongate 
rod-like in form. The oval or oat-shaped form is the common 
type in division rosettes. Division rosettes consisting of four 
small parasites found in smears from blood, spleen, and bone- 
marrow. Cross-form of division also met with. 


Fig. 156. — Babesia ninense (Yakimoff). (xc. 2500.) (After Sinton.) 

Dimensions. — Intracellular parasite, small form 0-7-1 /x, 
large form 2-3^ in length. 

Habitat. — Red blood- corpuscles in smears from the blood, 
Uver, spleen, and bone-marrow of the hedgehog, Erinaceus sp. : 
N.W.F. Province, Kohat. 

262. Babesia sergenti Wenyon. (Fig. 157.) 

Piroplasma ovis (part), Ratz, 1913, pp. 194—200. 
Theileria ovis, Yakimoff, 1916, p. 201. 

Oonderia ovis, Sergent, Parrot & Hilbert, 1922, pp. 789-92, 1 fig. 
Piroplasma ovis, Velu, 1922, pp. 216-23. 
Gonderia ovis, Lestoquard, 1924, pp. 122—8, 15 figs. 
Babesia sergenti, Wenyon, 1926, pp. 1004, 1007-8, fig. 414, 1-4. 
Babesia ovis, Reichenow, 1929, p. 1036. 
^Babesia sergenti, Krishna Iyer, 1933, p. 33. 

Morphologically the parasite corresponds very closely to 
B. mutans. Similar rounded and bacillary forms occur. 
Reproduction is by budding into two or four, so that the 
^characteristic cross-forms are produced. Nothing is known 
of the method of transmission. 


Remarks. — The piroplasms of sheep and goats continue to 
be referred to Piroplasma ovis Laveran & Nicolle (1909), but 
Wenyon (1926) has restricted that name to the organisms of 
intermediate size, and named the larger form as B. motasi 
and the smaller form as B. sergenti. Thus B. motasi, B. ovis, 

Fig. 157. — Babesia sergenti Wenyon. ( x 3000.) 
(From Wenyon, after Lestoquard.) 

and B. sergenti of sheep and goats correspond to B. bigemina, 
B. bovis, and B. mutans of cattle. B. sergenti produces no 
recognizable symptoms. 

Habitat. — Blood of goat, Capra hircus Linn. : United 
Provinces, Muktesar. 

263. Babesia soricis (Christophers). 

^Piroplasma soricis, Nuttall, 1908. 
Babesia soricis, Hoare, 1930, p. 246. 

Parasites of moderate size, occupying one-third of the cor- 
puscle in the larger forms. Did not show typical binary forms. 

Remarks. — ^NuttaU (1908) stated that " Christophers has 
found a parasite in musk rats in India which he names 
P[iroplasmM] soricis." Hoare (1930), faihng to trace the 
original pubhcation, referred to Christophers, who gave 
him the meagre information quoted above and remarked 
that he had found the parasite, but never described it or referred 
to it in print. 

Habitat. — Red blood-corpuscles of the musk rat, Crocidura 
cserulea (Kerr) : Madras, Guindy. 

264. Babesia tropicus (Lingard & Jennings). (Fig. 158.) 

"fPiroplasma tropicus (part), Lingard & Jennings, 1904, pp. 161-5 ; 
Baldrey, 1910, pp. 572-7, pi. xxxix. 

Small ring-forms, with central position of the chromatin in 
many of the infected red corpuscles. Chromatin in the form 
of two dots and a connecting bar between them. Pear-shaped 
forms rare in corpuscles, but occasionally seen free in the 

Remarks. — Lingard and Jennings (1904) found piroplasms 
in bovines, buffaloes, equines, elephants, camels, goats, sheep, 



dogs, cats, monkeys, deer, fowls, guinea-pigs, and lizards, 
without referring them to different species. Baldrey (1911) 
not only referred a form found in the blood of cavalry and 
artillery horses at Mhow and Meerut to this species, but also 
thought that B. gibsoni (Patton) was identical with it. Lingard 
and Jennings, and Baldrey were wrong in referring the parasite 
from cattle, dogs, sheep, and horses to P. tropicus when 
P. bovis, P. canis, P. ovis, and P. equi were aheady known from 
those animals. Since then other species have also been 
described from these animals, and piroplasms of monkeys 

Fig. 158. — Babesia tropicus (Lingard & Jennings). A, from camei ; 
B, from guinea-pig ; C, from elephant ; D, from the common 
hzard. (After Lingard and Jennings.) 

and deer have been described as P. pithed and P. cervi. 
No one else seems to have described piroplasms from elephants, 
camels, fowls, guinea-pigs, and hzards, and for this residue 
the name P. tropicus may be retained till the parasites from 
these animals are examined again and the specific name 
restricted to one particular form. 

Habitat. — Red blood- corpuscles of elephants, camels, guinea- 
pigs, fowls, and lizards : tJisriTEi> Pkovinces ; Bareilly. 


265. Babesia sp. 

■^Piroplasma sp., Patton, 1910, p. 279 ; 1911, p. 620. 
Babesia sp., Wenyon, 1926, p. 1027. 

Remarks. — The form may perhaps be the same as Babesia 
cervi Bettencourt, Franca and Borges (1907) described from 
Cervus dama Robert of Portugal. 

Habitat. — Blood of the spotted deer, Axis axis (Erxl.) 
(^Cervus axis) : India. 

266. Babesia sp. 

Babesia sp., Plimmer, 1915, p. 130 ; Wenyon, 1926, p. 1356. 

Remarks. — Wenyon (1926) thinks that this form may possibly 
be the same as B. gibsoni. He saw the parasite in large 
numbers in the blood of a Colombo dog. 

Habitat. — Blood of the wild dog, Cyon dukhunensis (Sykes), 
from India, in the Zoological Gardens, London ; Ceylon, 

267. Babesia sp. 

■\Piroplasma sp., Patton, 1910, p. 279 ; 1911, p. 620. 
Babesia sp., Wenyon, 1926, p. 1025. 

Remarks. — The form may possibly be the same as Babesia 
herpestidis Franca (1908) from the mongoose, Herpestes 
ichneumon Ilhger, in Portugal. 

Habitat. — Blood of Herpestes edwardii Desmarest [=Her- 
pestes mungo (Gmel.)]. 

Doubtful Protozoa. 

Incekt^ sedis. 

Family ANAPLASMIDiE Metz, Alexander & 
du Toit, 1933. 

Genus GRAHAMELLA Brumpt, 1911. 

{Syn. Bartonella Strong, Tyzzer, Brues, Sellards, and 

Gastiaburu, 1915). 

A new Hsematozoan, Graham -Smith, 1905, pp. 453-9, pis. xiii, xiv. 

Grahamella, Brumpt, 1911, pp. 514-17 ; 1913. 

Graham-Smith bodies, Laveran & Marullaz, 1914, pp. 240-6, pi. ii. 

■Orahaniella, Carini, 1915, pp. 103-4. 

Bartonella, Strong, Tyzzer, Brues, Sellards, & Gastiaburu, 1915, 

p. 32 ; Mayer, Borchardt, & Kikuth, 1926, p. 559 ; Wenyon, 

1926, pp. 1059-60 ; Noguchi & Battistini, 1926, p. 851 ; Noguchi, 

1926, pp. 533, 697, 715, 729. 
Grahamella, Wenyon, 1926, pp. 1056-9 ; Knowles, 1928, pp. 462-3 ; 

Brumpt, 1928, pp. 1079-81. 


Bartonella, McCamson& Mula Singh, 1931, pp. 945-9. 
Grahamia, Nietz, Alexander & du Toit, 1933, pp. 263-71. 
Bartonella, Donatien & Lestoquard, 1934, pp. 650-2 ; Adler & 

Ellenbogen, 1934, pp. 219-21 ; Kikuth, 1934, pp. 1241-50. 

Lestoquard, 1934, pp. 650-2 ; Adler & Ellenbogen, 1934, pp. 219- 

(Bartonella), Topley & Wilson, 1936, pp. 710-11. 
Qrahamella, Topley & Wilson, 1936, p. 712. 

Parasites minute ; rounded, oval or more usually rod- 
like bodies, showing an irregular staining, a blue cytoplasmic 
portion being distinguishable from a red chromatinic part. 
Rods may be straight or shghtly curved, and forms with 
a red- staining granule at each end with a constriction in the 
middle suggest that reproduction is by binary fission. One, 
two or as many as fifty may infect the same red corpuscle 
or an endothehal cell. 

Remarks. — Graham-Smith (1905) described these bodies in 
red blood- corpuscles of moles. Brumpt (1911), who en- 
countered the same structures in the red blood-corpuscles of 
moles in France, described them as a new genus. Other 
observers have found similar structures in the red blood - 
corpuscles of various other mammals. Wenyon (1926) 
thought that there was little evidence for regarding them as 
parasites, and still less that they were Protozoa. Structurally 
they resemble bacilli more than any other organisms. KJaowles 
(1928) supported this view. Reichenow (1929), Kudo (1931), 
and Calkins (1933) have left them out of consideration, 
probably because they do not believe them to be Protozoa. 

Very similar organisms were found in, human red blood- 
corpuscles and placed in a new genus called Bartonella. 
These are beheved to be the causal agents of two diseases, 
namely, Oroya fever and Verruga peruviana, occurring in 
man in Peru. The former is a fever with marked anaemia, 
and the latter a nodular eruption of the skin. Strong, Tyzzer, 
Brues, Sellard, and Gastiaburu (1915) estabHshed that these 
diseases are distinct, and gave the name Bartonella bacilliformis 
to the organism associated with Oroya fever. Noguchi and 
Battistini (1926) succeeded in cultivating the organism in vitro. 
They also succeeded in inoculating the form into Macacus 
rhesus (Audeb.). Noguchi in a series of papers (1926, 1927) 
has further shown that on subpassage in monkeys the virulence 
of the organism was increased, and it produced more marked 
anaemia than at first. Tb<^ organism was constantly present, 
as shown by culture, in tiie lymph glands, less often in the 
spleen, bone-marrow, and heart-blood. Inoculation of the 
monkey might produce either Oroya fever or Verruga peruviana, 
or both the conditions simultaneously. He also established by 
cross-immunity experiments that the two conditions were 
caused by the same virus. He further succeeded in transmitting 



the infection from monkey to monkey by the bites of the 
tick, Dermacentor andersoni. 

Mayer (1925) described another species of the genus in mice, 
and Donatien and Lestoquard (1934) yet another in cattle. 

Brumpt (1928) recalls in detail the successive creation of 
the genera Grahamella and Bartonella, and concludes that there 
is now insufficient cause for maintaining the genus Bartonella, 
created on the sole characteristic of pathogenicity. He 
regards Bartonella muris Mayer, 1921, as synonymous with 
Grahamella muris Carini, 1915, and combines under Grahamellu 
Brumpt, 1911, all forms described under the genus Bartonella, 
-which decision is followed here. 

Nietz, Alexander, and du Toit (1933), however, regard the 
two genera as distinct, and change the name Grahamella to 
Grahamia, but their paper is not available to the author. 
Topley and Wilson (1936) also regard them as distinct, and 
remark that the evidence in favour of Bartonella being a Uving 
reproducible micro-organism, capable of giving rise under 
favourable conditions to disease, is now very strong, but 
considerably less is known about Grahamella, and it would 
be unwise to assert at the moment that these bodies are 
definite bacteria. According to them, Grahamella, though 
resembling Bartonella, are much coarser, and more like 
ordinary bacteria. With Giemsa's stain they take on a blue 
rather than a reddish tint. Only occasional red cells are 
affected. Grahamella appear to be non-pathogenic and are 
not influenced by splenectomy. They have not so far been 
cultivated, and though it is probable that they are living 
■organisms, the evidence in favour of this is not yet conclusive. 

-268. Grahamella canis (Kikuth). 

Bartonella canis, Kikuth, 1928, p. 1729 ; 1929, pp. 1-7 ; Topley 

& Wilson, 1936, p. 711. 
■^Bartonella canis, Ray & Idnani, 1937, p. 371 ; 1938, p. 259. 

Forms in the red blood- corpuscles very pleomorphic, large 
or small, coccoid or rod-shaped. 

Remarks. — Kikuth (1928) described this form and con- 
sidered it responsible for causing infectious ansemia of dogs. 
He was unsuccessful in cultivating the organism on artificial 
media. Ray and Idnani (1937) have recently recorded the 
occurrence of this parasite in dogs in India, and beheve it 
to be the cause of one of the types of obscure canine fever. 
Appearance of the parasite in the blood was marked by 
a distinct thermal reaction associated with a number of 
pathological changes. Kikuth (1927) and others have stated 
that Bartonella usually appears in the blood of splenectomized 
•dogs, but Ray and Idnani (1938) find the infection to be 


inoculable into healthy non-splenectomized dogs, producing 
acute sjmaptoms and ultimate death of the host in several 

Habitat. — Blood of the dog, Ganis familiaris Linn. : United 
Provinces, Muktesar. 

269. Grahamella muris Carini. 

Grahamella anuria, Carini, 1915, pp. 103-4. 

Bartonella muris, Mayer, 1925 ; Mayer, Borchardt, & Kikuth, 
1926, p. 559 ; Wenyon, 1926, p. 1060. 
■fGrahatnella sp., Knowles, 1928, p. 463. 
^Bartonella muris, MeCarrison & Mula Singh, 1931, pp. 945—9. 

Dot- and rod-like bodies in the red blood- corpuscles of mice. 

Remarks. — Mayer (1925) described this organism from the 
red blood- corpuscles of mice which had been treated with 
Bayer 205 for trypanosome infections. Later, Mayer, Bor- 
chardt, and Kikuth (1926) showed that the same bodies appear 
in anaemia caused by splenectomy in rats. They believed 
that the operation had stimulated a latent infection. 
MeCarrison (1927) and WiUs (1930) showed that the same type 
of anaemia may arise in a proportion of non-splenectomized 
rats when they are fed on a diet deficient in fat- soluble vitamins 
and vitamin C. The faulty food would thus appear to induce 
functional injury to the spleen, thereby lowering resistance to 
infection, in a manner comparable to, though not so effective 
as, splenectomy. MeCarrison and Mula Singh (1931) have 
further shown that the blood of one- to four-day-old suckling 
rats born of splenectomized or non-splenectomized mothers 
may also show the infection. 

Knowles (1928) recorded Grahamella in certain films of rats' 
blood prepared by Parmanand of Bombay. Up to eight 
forms per red blood- corpuscle were encountered, while a very 
few lay apparently free and extra-corpuscular. 

Habitat. — Blood of rat : Madras, Madras ; Bombay, Bom- 

270. Grahamella sp. (Fig. 159.) 

" Corps Bartonelliformes," de Mello, 1928, pp. 515-16, fig. B, 29-38. 
Intra-corpuscular bodies, bipolar, navicular, fusiform or 


Fig. 159. — Grahamella sp. (After de Mello.) 



annular, the chromatin being concentrated into an excentric 
nucleus or at the two poles. 

Remarks. — De Mello regards these bodies of doubtful 
nature, and intermediate between Grahamella and Para- 

Habitat. — Blood of the ant-eater, Manis pentadactyla Linn. : 
Portuguese India, Nova Goa. 

Genus PARAPLASMA Seidelin, 1912. 

" Seidelin bodies," Seidelin, 1911, pp. 282-8. 

Paraplasma, Seidelin, 1912, pp. 503-4 ; 1914 a, pp. 203-8 ; Wenyon 

& Low, 1914, pp. 369-72 ; 1915, pp. 55-6 ; Seidelin, 1915, 

pp. 38-40 ; Seidelin & Connal, 1915, pp. 427-48. 
" Seidelin bodies," Thompson, 1915, pp. 479-82 ; Fowler, Simpson, 

Ross, Leishman, & Balfour, 1915, 3rd report ; Cropper & Drew, 

1916, pp. 20-4 ; de Mello, 1917, pp. 21-4. 

Polymorphic intra-corpuscular bodies, ranging from a small 
chromatic granule to large bodies consisting of a chromatic 
granule with a shaft of cytoplasm. 

Remarks. — SeideUn, who first described these bodies, con- 
sidered them to be the causative agent of yellow fever, and 
named the organism Paraplasma flavigenum. Seidelin and 
Connal (1915) considered the organism to be capable of trans- 
mission by artificial infection, and claimed to have discovered 
naturally infected guinea-pigs in West Africa, which were 
supposed to be reservoirs of the disease. Wenyon and Low 
(1914, 1915) refuted the parasitic view and showed that similar 
bodies were found in guinea-pigs in England, and especially in 
young animals. Finally, this latter view was supported by the 
Yellow Fever Commission, who reported (1915) that P. flavi- 
genum was not an organism, and was not connected with 
yellow fever. Cropper and Drew (1916) reported the occurrence 
of similar bodies in ansemic foetal blood. 

271. Paraplasma sp. (Fig. 160.) 

t" Seidelin bodies " (?), de Mello, de Sa, de Sousa, Dias, & Noronha, 
1917, pp. 21-4, pi. ii, figs. 1-20. 

Circular, vacuolated bodies occurring in groups of two, three 
or four in red blood-corpuscles, or in the form of thin or thick 
bacilli or vibrios. 


A B C D £ F 

Fig. 160. — Paraplasma sp. (After de Mello and others.) 


Remarks. — De Mello found these bodies in the blood-corpuscles 
of a man who had never been outside India, but it is doubtful 
if the so-called " Seidehn bodies " are an independent organism. 

Habitat. — Blood of man : Portugubsb India, Nova Goa. 

Genus ANAPLASMA Theiler, 1910. 

" Marginal points," Smith & Kilborne, 1893, p. 177. 

Anaplasma, Theiler, 1910, pp. 135-7 ; 1912 a, pp. 319-38 ; 1912 6, 
pp. 105-16; 1912 c, pp. 193-207; Minchin, 1912, p. 383; 
Dias & Aragao, 1914, pp. 231-49 ; Carpano, 1914, pp. 42-53 ; 
Lignieres, 1914, pp. 133-62; 1919 a, pp. 558-66; 1919 b, 
pp. 641-51 ; 1919 c, pp. 774-9 ; Hegner & Taliaferro, 1924, 
pp. 305-6 ; Wenyon, 1926, pp. 1053-6 ; Knowles, 1928, pp. 466-7. 

Spherical body of chromatin less than 0-5//, in diameter, 
situated at or near the edge or towards the centre of the red 
blood -corpuscles. Infection transmitted by a tick. 

Remarks. — Smith and KLilborne (1893) in their studies on 
Texas fever of cattle recognized certain red-staining granules 
on the margin of red corpuscles (" marginal points ") and 
regarded them as a resistant phase of Babesia bigemina. 
Theiler (1910) recognized similar bodies in the blood of cattle 
in South Africa, and referred them to Anaplasma, a new 
genus of parasitic Protozoa. The bodies are commonly found 
in association with infections of B. bigemina and B. mutans, 
and Carpano (1914) considered them as resistant forms of 
Piroplasmata. Dias and Aragao (1914) beheved that they are 
not true organisms at all, but a hsemolytic product. Theiler 
(1912) and Lignieres (1919) have described cases of pure 
infection with these bodies. According to the latter author, 
in true Anaplasma infections as many as 50 per cent, of the 
red corpuscles may be infected, and the infection is readily 
transmissible to clean cattle. Infected cattle suffer from 
fever and an intense and progressive anaemia. Laveran and 
Franchini (1914) discovered them in numerous Mammals, 
such as rats, mice, rabbits, guinea-pigs, moles, cats, dogs, 
calves, pigs, and monkeys. De Mello and his colleagues (1917) 
found them in human blood, and also in the blood of fishes, 
frogs, lizards, snakes, and birds, but did not regard them as 
Protozoa, considering them to be hsemolytic products. 

272. Anasplasma sp. (Fig. 161.) 

■\ Anaplasma, de Mello, de Sa, de Sousa, Dias, & Noronha, 1917, 
pp. 17-21, pi. ii, figs. 21-9. 

Intra-corpuscular, oval or lanceolate, of small dimensions, 
and staining like chromatin ; apparently not possessing 
any cytoplasm. 

326 . SPOROZOA. 

Bemarks. — De Mello and his colleagues (1917) found 
Anaplasmosis in the blood of fishes, frogs, lizards, snakes, 
tortoises, and man. Discussing the origin of the so-caUed 
Anaplasma, they came to the conclusion that they are not 
Protozoa, but are unquestionably due to hsemolytic altera- 
tions. They regarded them as due to nuclear fragmentation,. 


-Anaplasma sp. (After de Mello and others.) 

which is normal in Vertebrates with nucleated red corpuscles, 
or pathological, as in anaemias of Mammals. 

Habitat. — Blood ofPercafluviatilis Ijinn.,Eana tigrina Daud., 
Eana esculenta Linn., Hemidactylus brooki Gray, Coluber 
blumenbachii Schlegel, Naja naja Linn., Ghitra indica (Gray), 
and Homo sapiens Erxleben : Portuguese India, Nova Goa. 

273. Anaplasma sp. 

'\Anaplas'ma sp., Knowles, 1928, p. 467. 
Habitat. — Blood of Indian bats : Bengal, Calcutta. 

Genus BERTARELLIA Carini, 1930. 

Bertarellia, Carini, 1930, pp. 1312-13 ; de Mello & de Meyrelles, 
1937 b, pp. 98-108. 

Polymorphic intra-corpuscular bodies, probably of a para- 
sitic nature ; usually round or oval, variable in size, smallest 
of the size of cocci and the largest 1-2 ^u, in diameter. Stained 
with Leishman's or Giemsa's stains, in the largest individuals 
the cytoplasm is stained blue, usually with a chromatic granule 
stained red. The infected red blood- corpuscles are not altered. 

274. Bertarellia calotis de MeUo & de Meyrelles. (Fig. 162.) 

■fBertarellia calotis, de Mello & de Meyrelles, 1937 b, pp. 98-108, 
pi. xii, 2 text-figs. 

Circular bodies, with refringent greenish cytoplasm 
surrounded by a strong membrane, intra-corpuscular or free, 
dividing by a process of budding. Stained by Leishman's 
or Giemsa's stains they show a variety of forms ; they appear 
either as a small round chromatic dot, of anaplasmoid nature. 


or like a vacuole devoid of any central granule or a small 
roundish body surrounded by a more or less strong membrane, 
taking a chromatic stain, with a central nuclear granule. 
The cytoplasm is stained blue or greyish-blue, often surrounded 
by a white circular halo, without any granule at all or with 
a central chromatic granule or the nuclear mass may be 
located at the periphery attached to the membrane. The 
infected red blood- corpuscle may have two or three of these 
parasites, and does not show any alteration. 

A B C D E F G 

Fig. 162. — Bertarellia calotis de Mello & de Meyrelles. A, anaplasmoid ; 
B, surrounded by a vacuole ; C, cytoplasm surrounded by 
a strong membrane ; D, cytoplasm containing a central 
chromatic granule ; E, cytoplasm with peripheral chromatic 
mass ; F, with central and peripheral chromatic granules ; 
G, chromatic granule showing budding. (After de Mello and 
de Meyrelles.) 

Remarks. — Blood-smears fixed with Bouin's or Schaudinn's 
fluid and stained with Heidenhain's iron-hsematoxylin are 
said to have confirmed the structure as revealed by Romanow- 
sky's stain, and have thrown fight on the process of division. 
The chromatic granule or nucleus becomes enlarged and takes 
a ring form. Later it becomes compact and gives origin 
to a small bud, which protrudes to the exterior, becomes 
covered by membrane and separated from the main ceU. 

Habitat. — Blood- corpuscles of Galotes versicolor Daud. 
subspecies major Blyth : Portuguese India, Nova Goa. 

275. Bertarellia sp. 

^Bertarellia sp., de Mello & de Meyrelles, 1937 b, p. 107. 

Habitat. — Blood of the Indian tortoise, Lissemys punctata 
granosa (SchoepfiF) : Portuguese India, Nova Goa. 



II. Subclass CNIDOSPORIDIA Doflein, 

This subclass includes organisms which are amoeboid during 
the trophic phase, and of which dissemination takes place 
through resistant spores, which are provided with one to four 
polar capsules. Each spore contains one to many sporoplasms 
or generative cells, and the spore -membrane may be complete 


Fig. 163.^ — Some typical Cnidospobidia and their spores. A, Chloro- 
myxum leydigi Mingazzini ; B, C, Leptotheca agilis Thelohan 
in different stages of locomotion ; D, spore of the same ; 
E, spore of Henneguya psorospermica Thelohan, stained 
section, ect., ectoplasm ; end., endoplasm ; f.g., fat- 
globules ; n., nuclei ; p.c, polar capsules ; ps., pseudo- 
podia ; s., sutural plane ; sp., spores ;, stylar 
pseudopodium ; v., vacuole. (From Reichenow, after 
Doflein and Thelohan.) 

or be bivalved or trivalved. Each polar capsule contains 
a long coiled filament, which, when extruded, serves to attach 
the spore to the intestinal wall till the amoeboid body can 
escape from the spore and infect the tissues of the new host. 

Schaudinn (1900) divided the Sporozoa into two sub- 
classes, the Telosporidia and the Neosporidia. The Telo- 
SPORIDIA include the Gregarinida, Coccidia, and H^mo- 
SPORIDIA, and are a uniform group, characterized by having 


definite intracellular stages, schizogony, and, following upon 
conjugation of gametes, formation of oocysts within which the 
sporozoites are formed. The Neosporidia, including the 
Cnidosporidia, Sarcosporidia, and Haplosporidia, do not 
form a natural group, nor have they much in common with the 
Telosporidia. In the Neosporidia the life of an individual 
does not come to an end when reproduction takes place, but 
reproduction continues throughout the trophic phase, the 
sporoblasts being carried about by the more or less active 
organism, which may ultimately become a large mass of 
spores. Among the Cnidosporidia the parasites may some- 
times be intracellular, but they usually reproduce by binary 
fission, and the zygotes do not become encysted and do not 
produce sporozoites. The Sarcosporidia and Haplosporidia 
possess simple spores, and do not seem to be related either to 
the Cnidosporidia or to the Telosporidia. 

Wenyon (1926) considers it justifiable to reserve the title 
Sporozoa for the Telosporidia, to consider the Cnidosporidia 
as an independent class, and to regard Sarcosporidia and 
Haplosporidia as parasites of undetermined position. 
Reichenow (1929, 1935) regards Telosporidia, Cnido- 
sporidia, Sarcosporidia, and Haplosporidia as independent 
subclasses of Sporozoa. Kudo (1931) also follows this 
arrangement, but places Sarcosporidia and Haplosporidia 
in one subclass with the title Acnidosporidia. Calkins (1933) 
regards Telosporidia, Cnidosporidia, and Acnidosporidia 
as classes of the subphylum Sporozoa. 

Cnidosporidia are exclusively parasites of the Inverte- 
brates and the lower Vertebrates, and are responsible for 
causing epidemics of infection among animals of economic 
importance such as fishes, silkworms, and honey-bees. There 
are no secondary or intermediate hosts. 

Following Kudo, the Cnidosporidia are divided into four 
orders, as follows : — 

1. Spore large; membrane bivalved ; two or [Biitschli, p. 330. 

four polar capsules visible in vivo Myxosporidia 

2. Spore large ; membrane trivalved ; three dis- [Stole, p. 346. 

tinctly visible polar capsules Actinomyxidia 

3. Spore small ; raembrane in one piece ; one [Balbiani, p. 346. 

(rarely two) polar filaments ; invisible in vivo . Microsporidia 

4. Spore small, barrel-shaped ; a thick filament 

coiled beneath the spore-membrane ; three [Kudo, p. 360. 

sporoplasms Helicosporidia 



I. Order MYXOSPORIDIA Biitschli, 


The Myxosporidia are distinguished by their characteristic 
spores, which are of various shapes and dimensions. Each 
spore is covered by a bivalved chitinous membrane called the 
spore -membrane, the two valves of which are united in a 
sutural plane, which may be more or less straight or irregularly 

E F G 

Fig. 164. — Some characteristic forms of spores of various Myxosporidia. 
A, Ceratomyxa truncata Thelohan ; B, Ghloromyxum 
■ leydigi Mingazzini ; C, Sphserospora rostrata Thelohan ; 
D, Myxidium procerum Auerbach ; E, Zschokkella acheilo- 
gnathi Kudo ; F, Myxoholus carassii Klokacewa ; G, Henne- 
guya gurleyi Kudo. ( X 1500.) (After Kudo.) 

curved . The surface of the valves may be smooth or marked 
with ridges, and the form of the spore depends upon the shape 
of the valves and the presence of accessory appendages. 

Within the shell are polar capsules, which may be one, 
two or four in number and are usually situated at what is 
described as the anterior end of the spore. In the family 
Myxidiidse there are two polar capsules, one near each pole 
of the spore. Each polar capsule contains a coiled filament, 
which can be extruded through its pore. The substance 
contained in the spore, apart from the polar capsules, is 
designated the sporoplasm. It usually contains two nuclei. 



and in the family Myxobolidse an iodinophilous vacuole 
containing glycogenous substance which stains mahogany- 
red with iodine. 

When introduced into the digestive tract of a fish, the 
sporoplasm leaves the spore as amoeboid " planonts " or 
wanderers. These probably fuse together and grow into the 
characteristic trophic phase, the so-called multinucleate 
Plasmodium. Passing through the epithelium of the gut, 
it enters the tissues of specified organs, grows into a schizont, 
and its nucleus divides repeatedly. Some of the nuclei 
become surrounded by cytoplasm and form sporonts. The 
sporont may be monosporous, disporous or polysporous, 
according as it produces one, two or many spores. 

Fig. 165.- — Development of the spore from the pansporoblast inMyxobolus 
pfeifferi Thelohan. A, single propagative cell from multi- 
nucleate Plasmodium ; B, division to form one large and 
one small cell ; G, association of two pairs to form a group 
of two large and two small cells ; D, formation of six- 
celled stage ; E, stage with fourteen nuclei, two of which 
are the nuclei of the original small cells ; F, division into 
two bodies, each with six nuclei, while the nuclei of the small 
cells take up a position at the angles between them ; 
G, each sporoblast now divides into three cells, two of which, 
each with a single nucleus and a vacuole, are the capsulo- 
genoiis cells containing the polar capsules, one with two 
nuclei forms the binucleate amcebula, while the remaining 
two nuclei become peripherally arranged and form, together 
with some cytoplasm, the valves of the spore ; H, more 
advanced stage of one of the developing spores ; /, J fully 
developed spores. (From Wenyon, after Keysselitz.) 



Sporulation begins with a peculiar process of endogenous 
or internal budding. An island of protoplasm or pansporo- 
blast is formed round two of the nuclei, which are usually 
dimorphic. Each of these nuclei undergoes division until 
fourteen are present, seven from each of the original nuclei. 
The bud then divides into two cells, each of which is a sporo- 
blast containing six nuclei, after one has been extruded. 
Two of these six nuclei form the valves of the capsule (sporo- 
cyst), two form the polar capsules, and two remain as pro- 
nuclei, which subsequently fuse when the spore becomes 
mature. Thus the endogenous bud represents a zygote, 
and the two original nuclei of the pansporoblast the pro- 
gametic nuclei. The details of the process of sporulation 


Fig, 166. — External appearance of Myxosporidian infection in jSsh. 
A, head of the short-headed red-horse, showing the cysts 
of Myxobolus conspicuus Kudo, x J ; -B, the river chub, 
with cysts of Myxobolus squamosus Kudo, X f ; C, a blunt- 
nosed minnow, with numerous cysts of Myxobolus aureatus 
Ward on the fins, x f . (After Kudo.) 

differ in different species. Neville (1931) has reviewed the 
previous literature and made an independent study of five 
different species. He concludes that there are two types of 
nuclei, germmal and vegetative. The germinal nuclei contain 
the diploid number of chromosomes, but after several divisions 
undergo reduction, whereby the number of chromosomes is 
reduced to one-half. 

The simpler members of the order occur in various cavities 
in the bodies of their host, such as the gall-bladder, uriniferous 


tubules of the kidney or urinary bladder, and do not cause 
much harm to the host. In other cases the parasites invade 
the tissues, causing them to degenerate, and give rise to 
tumours in muscles, central nervous system, etc., which are 
referred to as Myxosporidian cysts and are visible to the naked 
eye, or the parasites may spread through tissue, giving rise to 
diffuse infiltration. 

TheMYXOSPORiDiA are parasites of cold-blooded Vertebrates, 
more particularly fishes. Each species attacks one or several 
species of fish. In the case of freshwater fishes they usually 
invade the gills and the gall-bladder, while in the marine 
fishes they are usually found in the gall-bladder or the urinary 
bladder. Whitish pustules visible to the naked eye may 
indicate the infection in the giUs, and in the case of internal 
organs there may be abnormal changes of form or colour, 
but ordinarily infection can only be detected by microscopical 

Doflein (1901) divided the order into two suborders, Dis- 
POREA and PoLYSPOREA, but Kudo (1920) considers such 
a division an artificial one, as the number of spores produced 
by any species is not always constant. 

Following Kudo (1920, 1933) the Myxosporidia are divided 
into three suborders, as follows . — 

1. Largest diameter of the spore at right angles to the 

sutural plane ; one polar capsule on each side of 

the plane ; sporoplasm without iodinophilous [Kudo, p. 333. 

vacuole Eurysporea, 

2. Spore spherical or subspherical, with one, two 

or four polar capsules ; sporoplasm without [Kudo, p. 335. 
iodinophilous vacuole SphaBFOSporea, 

3. Sutm-al plane of the spore coincides or forms 

an acute angle with the longest diameter ; one 

or two polar capsules ; sporoplasm with or [Kudo, p. 336. 

without an iodinophilous vacuole Platysporea, 

I. Suborder EURYSPOREA Kudo, 1920. 

The largest diameter of the spore is at right angles to the 
sutural plane, with one polar capsule on each side of the plane. 
Sporoplasm without iodinophilous vacuole. Vegetative form 
found in body- cavity. The great majority are parasites of 
marine Fish. Monosporous, disporous or polysporous. 

Identification Table of Families. 

i. Typically coelozoic, in marine Fish . . Ceratomyxidse Doflein, p. 334. 
2. Histozoic or coelozoic, in fresh- 
water Fish Wardiidse * Kudo. 


Family CERATOMYXID^ Doflein, 1899. 
Typically coelozoic parasites of marine Fish. 

Genus CERATOMYXA Thelohan, 1892, emend. 

Ceratomyxa, Thelohan, 1892, pp. 169, 171, 175 ; Labbe, 1899, 
pp. 89-91 ; Minchin, 1903, pp. 289, 293, 295, 298 ; 1912, p. 408 ; 
Kudo, 1920, pp. 61-80 ; Wenyon, 1926, p. 725 ; Eoiowles, 1928, 
p. 324; Reichenow, 1929, pp. 195.3-7; Kudo, 1931, p. 306; 
1933, pp. 196, 199-201 ; Calkins, 1933. p. 567 ; Reichenow, 1935, 
p. 386. 

Spore arched, breadth more than twice the sutural diameter. 
Valves of spore extended into long, lateral, conical, and 
hollow processes. Sporoplasm asymmetrically placed, usually 
not filling the intrasporal cavity. Monosporous, disporous 
or polysporous. Generally in the gall-bladder or urinary 
bladder of marine Fish. 

Key to Indian Species. 

1(2). Spore crescent-shaped, 14-15 /i by 4r-5;Lt. C . gohioidesi Chak.., 

[p. 334. 
2 (1). Spore elliptical, 30-40 ja by lO/ii C. hilsse Chak., p. 335. 

276. Ceratomyxa gobioidesi Chakravarty. 

^Ceratomyxa sp., Ray, 1933 a, p. 259 ; 1933 b, p. 349. 
^Ceratomyxa gobioidesi, Chakravarty, 1938. 

Trophozoites more or less spherical, not showing sharp 
demarcation between ectoplasm and endoplasm, pseudo- 
podia short and blunt. Disporous ; spores crescent-shaped, 
valves sjnnmetrical and terminating in blunt points, sutm-al 
plane distinct. Extra-capsular cavity filled with finely 
granular sporoplasm. Polar capsules equal and spherical, 
situated on each side of the sutural plane, and provided with 
distinct coiled filaments. 

Dimensions. — Trophozoites 500-650 /n in diameter ; spores 
14-15 )Li in breadth, 4-5 /x in sutural diameter ; polar capsules 
2-5-3 /A ; polar filament 15 /x. 

Remarks. — Ray (1933 a, b) reported this species from a 
number of estuarine fish, but did not publish any description. 
Chakravarti (1938) has recently restudied the form and 
described it as a new species. He has kindly allowed me to 
see the manuscript of his paper prior to its pubHcation, and 
to abstract the description of this and several other species. 

Habitat. — Liver, gall-bladder, kidney, ovary, etc., of 
Gobioides rubicundus Hamilton ; gall-bladder of Tricho- 
gaster fasciatus Schneider and Macrones gulio (Ham.-Buch.) : 
Bengal, Calcutta. 


277. Ceratomyxa hilsae Chakravarty. 

"fCertaomyxa hilsas, Chakravarty, 1938. 

Trophozoites in large numbers in the gall-bladder of the 
host. Disporous, spores often seen lying side by side in a com- 
mon envelope. Spores elliptical, valves tapering to blunt 
ends, sutural plane prominent, diAdding the spore into equal 
parts. Extra-capsular cavity filled with finely granular 
sporoplasm. Polar capsules spherical, of equal sixe, provided 
with well-marked coiled polar filaments. 

Dimensions. — Spores 30-40 ju, in breadth, 10 /x in sutural 
diameter ; polar capsules S^u. in diameter ; polar filament 
35^0 fx. 

Bemarks. — The above description is based on Chakravarty's 
paper in manuscript. 

Habitat. — Gall-bladder of Hilsa ilisha (Ham.-Buch.) : 
Bengal, Calcutta. 

II. Suborder SPH^ROSPOREA Kudo, 1920, 

Spore spherical, with one, two or four polar capsules. 
Sporoplasm without iodinophilous vacuole. Vegetative form 
found in body-cavity and tissues. Monosporous, disporous 
or polysporous. Parasites of marine and freshwater Fish 
and Amphibia. 

Identification Table of Families. 

1. With foTir polar capsules ChloromyxidSB Thelohan, 

[p. 335. 

2. With two polar capsules Sphserosporidse Davis, 

[p. 336. 

3. With one polar capsule Unicapsulidse * Kudo. 

1. Family CHLOROMYXIDiE Thelohan, 1892. 

Spore with four polar capsules. Monosporous, disporous 
or polysporous. 

Genus CHLOROMYXUM Mingazzini, 1890. 

Chloromyxum, Mingazzini, 1890, p. 160 ; Labbe, 1899, pp. 94-6 ; 
Minchin, 1903, p. 295 ; 1912, p. 409 ; Kudo, 1920, pp. 87-99 ; 
Wenyon, 1926, p. 726; Reichenow, 1929, pp. 1058-9; Kudo, 
1931, p. 308 ; 1933, pp. 197, 202-3 ; Calkins, 1933, p. 568 ; 
Ray, 1933, p. 259 ; Reichenow, 1935, p. 386. 

Spore with four polar capsules, grouped at the anterior end ; 
surface often striated or with ridges ; sutural Une often obscure. 
Monosporous, disporous or polysporous. Histozoic, or coelo- 
zoic in freshwater and marine Fish and also in Amphibians. 


278. Chloromyxum amphipnovi Ray. 

fChloromyxum amphipnovi, Ray, 1933 a, p. 259. 
f Chloromyxum sp., Ray, 1933 b, p. 349. 

Tiiis parasite was reported by Ray (1933) as a new species, 
but no account of it has as yet been pubJished. 

Habitat — Gall-bladder of Amphinous kuchia (Ham.-Buch ) : 
Bengal, Calcutta 

2. Family SPH^ROSPORID^, Davis, 1917. 

Spore with two polar capsules. Monosporous, disporous 
or polysporous. 

Genus SPH^ROSPORA Thelohan, 1892. 

Spheerospora, Thelohan, 1892, p. 167 ; Labbe, 1899, p. 86 ; Minchin, 
1903, pp. 286, 293, 295, 298 ; Kudo, 1920, pp. 100-4 ; Wenyon, 
1926, p. 726 ; Reichenow, 1929, p. 1059 ; Kudo, 1931, p. 308 ; 
1933, pp. 197, 203 ; Calkins, 1933, p. 568 ; Reichenow, 1935, 
p. 387. 

Spore with two polar capsules at anterior end. Sutural line 
straight. Monosporous, disporous, or polysporous. Body- 
cavity and tissue parasites of freshwater and marine Fish. 

279. Sphserospora sp. 

■\ Spheerospora sp., Southwell & Prashad, 1918, pp. 347-8. 
Sphserospora sp.. Kudo, 1920, p. 103 ; 1933, p. 203. 

Remarks. — The poor condition of the material did not allow 
of a complete account of its structure being given, but the 
bicapsulate, rounded structure of the spore places it in this 

Habitat. — Cysts occurred in very large numbers, one under 
each scale of Barilius barna (Ham.-Buch.), from the vicinity 
of the Ruby Mines : Burma. 

III. Suborder PLATYSPOREA Kudo, 1920. 

Sutural plane coincides or forms an acute angle with the 
longest diameter. One, two or four polar capsules. Sporo- 
plasm with or without an iodinophilous vacuole. 

Identification Table of Families. 

1 (5). Without iodinophilous vacuole . 2-4. 

2. Two polar capsules, one at each 

pole Myxidiidae Thelohan, p. 337. 

3. One polar capsule Coccomyxidse * Leger & Hesse. 

4. Two or four polar capsules at 

anterior end Myxosomatidse * Poche. 

5(1). With an iodinophilous vacuole . Myxobolidse Thelohan, p. 340. 


1. Family MYXIDIIDiE Thelohan, 1892. 

Spore fusiform or semicircular. One polar capsule at each 
end. Sporoplasm without iodinophilous vacuole. 

Key to Indian Genera. 

1. Polar filaments long and fine 2. 

2 (3). Spores fusiform, with pointed ends ; [p. 337. 

polar capsules oppositely directed . . . Myxidium Biitschli, 

3 (2). Spores fusiform, usually with truncated [bach, p. 339. 

ends ; polar capsules obliquely directed. Zschokkella Auer- 

Genus MYXIDIUM Biitschh, 1882. 

Myxidium, Biitschli, 1882, p. 593. 
Cystodiscus, Tuntz, 1889, pp. 84-8. 
Myxidium, Labbe, 1899, pp. 91-2 ; Minchin, 1903, pp. 286, 288, 

295 ; 1912, p. 409 ; Kudo, 1920, pp. 108-18 ; Wenyon, 1926, 

p. 726 ; Reichenow, 1929, pp. 1060-2 ; Kudo, 1931, p. 309 ; 

1933, pp. 197, 204-6 ; Calkins, 1933, p. 568 ; Reichenow, 1935, 

p. 387. 

Spore fusiform, with pointed or rounded ends. Polar 
capsules typically pyriform. Polar filament comparatively 
long and fine. Monosporous, disporous or polysporous. 
Typically coelozoic, but also histozoic in marine or freshwater 
Pish, Amphibians or Reptiles. 

Key to Indian Species. 

1 (2). Spore elongated, fusiform, 10-12 /i in [p. 337. 

length M. danilewskyi Lav., 

2(1). Spore not fusiform 3. 

3 (4). Spore oval, tapering at each end to [p. 338. 

a blunt point, 16/n by 5/x M. mackiei Bosan., 

4 (3). Spore ovoidal, 12-15fi by 8-5-10/x .... M. glossogobii Chak., 

[p. 337. 

280. Myxidium danilewskyi Laveran. 

Myxidium danilewskyi, Laveran, 1897, p. 725 ; 1898, pp. 27-30, 
30 figs. ; Labbe, 1899, p. 92. 
•f Myxidium danilewskyi, Laveran & Mesnil, 1902, p. 609. 
Myxidium danilewskyi, Minchin, 1903, pp. 275, 294, 346 ; Kudo, 
1920, pp. 109-10, figs. 255-7 ; 1933, p. 204 ; Reichenow, 1935, 
p. 387. 

Vegetative phase not known . Disporous . Spores elongated, 
fusiform, sometimes a Httle incurved. Polar capsules small, 
situated at the extremities. 

Dimensions. — Length of spore 10 jit. 

Habitat. — Prom the kidney of the tortoise, Chinemys 
reevesi (Gray) : Ceylon. 

281. Myxidium glossogobii Chakravarty. 

■\ Myxidium glossogobii, Chakravarty, 1938. 
Vegetative form not observed. Spores elongate oval, 
SPOR. z 



valves not striated. Space outside the capsule filled entirely 
with uniformly granular sporoplasm, containing several black 
dots in the fresh condition. Polar capsules one at each end 
of the spore, spherical at first, but ovoidal after the extrusion 
of the polar filament. Openings through which filaments 
are extruded are marked by elevated areas of the shell just 
in front of the polar capsules. 

Dimensions. — Spore 12-15 /i in length, 8-5-10 /x in breadth ; 
polar capsules S-l^-lfx, ; polar filament 40-50 /u.. 

Remarks. — Description is based on Chakravarty's paper in 

Habitat. — Gall-bladder of Glossogobius giuris (Ham.-Buch.) : 
Bengal, Calcutta. 

282. Myxidium mackiei Bosanquet. (Fig. 167.) 

fMyxidium m.ackiei, Bosanquet, 1910, pp. 436-8, figs. 7-13. 

The vegetative phase does not ordinarily exhibit a distinction 
between ectoplasm and endoplasm ; in a few cases where the 

Fig. 167. — Myxidium mackiei Bosanquet. A, vegetative phase in 
contact with cells of a renal tubule ; B, spores. (After 

whole organism was converted into spores there was a cuticle 
or cyst-wall. Cytoplasm contains a large number of nuclei, 
apparently of two varieties, some vesicular, others smaller 
and compact. Spores formed in pairs, at many points simul- 
taneously, ultimately the whole mass being converted into 
spores. A portion of the cytoplasm becomes rounded oflF 
and lies in a definite space Avithin the parasite. This portion 


(disporoblast ?) contained in one instance 10 nuclei, but 
it seems likely that 12 is the full number of nuclei in the 
disporoblast, as the final sporoblasts into which this divides 
appear to contain usually 6 nuclei. Of the 6 nuclei of the 
sporoblast 2 go to form the valves of the sheath, 2 attach 
themselves to the polar capsules, and 2 remain in the sporo- 
plasm. The two last-mentioned nuclei may fuse into one. 
In the sporoplasm two large vacuoles are often seen, one near 
each polar capsule. Both ends of a spore taper to a blunt 
point and the spore-coat is markedly striated. 

Dimensions. — Spores average 16 ju, in length by 5 ju, in breadth. 

Remarks. — ^The description is based on sections of the kidney 
of the host prepared by J. Percival Mackie in the Bombay 
Bacteriological Laboratory. 

Habitat. — Kjdney of the tortoise, Trionyx gangeticus Cuvier : 
Bombay, Bombay. 

283. Myxidium sp. 

•fMyxidium sp., Ray, 1933 b, p. 349. 

Habitat. — Gall-bladder ofClarias batrachus Cuvier & Valenc, 
Saccobranchus fossilis (Bloch.) ; Ophiocephalus punctatus 
Bloch : Bengal, Calcutta ; gall-bladder of Kachuga smithi 
(Gray), Lissemys punctata granosa (Schoepff) : Unitei> 
Provinces, AUahabad, and Geomyda trijuga (Schweigger) : 
Madras, Madras. 

Genus ZSCHOKKELLA Auerbach, 1910. 

Zschohkella, Auerbach, 1910; Kudo, 1920, pp. 122-3; Wenyon, 
1926, p. 727 ; Reichenow, 1929, p. 1062 ; Kudo, 1931, p. 310 ; 
1933, pp. 198, 206 ; Calkins, 1933, p. 568 ; Reichenow, 1935, 
p. 387. 

Spore semicircular in front view ; ellipsoidal in profile ; 
ends pointed. Sutural line much curved. Polar capsules 
large and spherical ; polar filaments long and fine. Mono- 
sporous, disporous or polysporous. Typically coelozoic 
parasites of fresh- and salt-water Fish. 

284. Zschokkella prashadi Ray. 

^Zschokkella prashadi, Ray, 1933 a, p. 259. 
fGystodiscus (Zschokkella) sp., Ray, 1933 b, p. 349. 

This parasite was reported by Ray (1933) as a new species, 
but no description has so far been pubHshed. 

Habitat. — Gall-bladder of Bufo melanostictus Schneid., 
Rana tigrina Daud. : Bengal, Calcutta ; also gall-bladder of 
Lissemys punctata granosa (Schoepff) : United Provinces, 



2. FamUyMYXOBOLID^Thelohan, 1892. 

Spore with one or two polar capsules at the anterior end, 
with or without posterior processes. Sporoplasm with an 
iodinophilous vacuole. Majority polysporous in freshwater 

Key to Indian Genera. 

1 (2). Each valve of spore prolonged into a [p. 344. 

long process Henneguya Thelohan, 

2(1). Valves without posterior processes. ... 3. [p. 340. 

3 (4). Spores ovoidal ; two polar capsules . . Myxobolus Biitschli, 

4 (3). Spores pyriform ; one polar capsule . . Thelohanellus Kudo, 

[p. 342. 

Genus MXYOBOLUS BiitschH, 1882. 

Myxobolus, Biitschli. 1882, pi. xxxviii, figs. 6-10 ; Labbe, 1899, 
pp. 97-101 ; Kudo, 1920, pp. 58, 128-58 ; Wenyon, 1926, p. 727 ; 
Reichenow, 1929, pp. 1668-72; Kudo, 1931, p. 312; 1933, 
pp. 198, 208-10 ; Calkins, 1933, p. 568 ; Reichenow, 1935, p. 388. 

Spore ovoidal, flattened. Shell without posterior processes. 
Two pyriform polar capsules at the anterior end. Sporo- 
plasm with an iodinophilous vacuole. This genus includes 
many species ; the majority are polysporous and tissue- 
parasites of freshwater Fish, but a few are known from marine 
Fish, an Annelid, and an Amphibian. 

Key to Indian Species. 

1 (4). Spore with unequal polar capsules 2. p. 340. 

2 (3). Spores oval, 12-4-15)iiby 8-2-lOitt ■ M. calbasui Chak., 

3 (2). Spores spherical to oval, 7-21-8-24/i, in [p. 341. 

length M. mrigalae Chak., 

4 (1). Spore with equal polar capsules 5. [& Prash., p. 341. 

5 (6). Spores ovoidal, 9/x by 7-2 /^ M. nodularis Southw. 

6 (5). Spores spherical, 13/n in diameter .... M. sp., Southw., 

[p. 342. 

285. Myxobolus calbasui Chakravarty. 

■fMyxoboltis sp. (part), Ray, 1933 6, p. 349. 
^Myxobolus calbasui, Chakravarty, 1938. 

Trophozoites spherical to oval. Polysporous. Spores oval, 
rounded at the posterior and pointed at the anterior end, 
sutural plane distinct. Sporoplasm occupjdng the posterior 
region of the spore. Polar capsules pyriform, one smaller 
than the other. Coiled polar filaments well marked, unequal 
in length when extruded. A spherical iodinophilous vacuole 

Dimensions. — Spore 12-4-15 ;u, in length, 8-2-10 /a in breadth ; 
polar capsules 6-18)u, by 4-12^ and 4-12/i, by 3"09yLt ; polar 
filaments 125 /x and 60 ft respectively ; iodinophilous vacuole 
4- 1 ju. in diameter. 

Remarks. — Description is based on Chakravarty's paper 
in manuscript. 


Habitat. — Ovary and liver of Glarias hatrachus Cuvier & 
Valenc. ; gills of Katla katla (Ham.-Buch.) ; liver of Girrhina 
mrigala (Ham.-Buch.) ; and gall-bladder of Labeo calhasu 
(Ham.-Buch.), Labeo rohita (Ham.-Buch.), and Girrhina 
mrigala (Ham.-Buch.) : Bengal, Calcutta. 

286. Myxobolus mrigalae Chakravarty. 

■fMyxobolus mrigalss, Chakravarty, 1938. 

Cysts oval, on the scales of the host, perforated, and con- 
taining large number of mature spores. Polysporous. Spores 
spherical to oval, valves thick, exhibiting several triangular 
markings in front view. Sporoplasm situated at the posterior 
end of the capsules. Polar capsules pyriform, unequal in 
size, divergent, and containing distinct coiled filaments. 
An iodinophilous vacuole is present. 

Dimensions. — Cysts *75-l-5 mm. in length and -75-1 mm. 
in breadth ; spores 7-2 1-8-24 ju, ; polar capsules 5-15/x by 
3-09/1 and 3-09 /x by 2-06 /x ; polar filaments 41-2 /x and 20-6 /i 

Remarks. — Description is based on Chakravarty's paper in 

Habitat. — Scales oi Girrhina mrigala (Ham.-Buch.) : Bengal, 

287. Myxobolus nodularis Southwell & Prashad. (Fig. 168.) 

^Myxobolus nodularis, Southwell & Prashad, 1918, p. 347, pi. xi, 
figs. 32-6. 
Myxobolus nodularis, Kudo, 1920, p. 153 ; 1933, p. 209. 

Cyst rounded or sHghtly elongated, creamy -yeUow in colour. 
Spore ovoidal. Suture between valves thick. Two polar 
capsules. Polar filament very much coiled. 

Fig. 168. — Spore of Myxobolus nodularis Southwell & Prashad. 
(After Southwell and Prashad.) 

Dimensions. — Cysts from 3-5 to 3-8 mm. in length and 
2-3-2-8 mm. in breadth ; spores 9/x by 7-2/x ; polar capsules 
3-4)u, in length, filament 18-3/^ in the extruded condition. 

Habitat. — ^Muscles of Rasbora daniconius (Ham.-Buch.) : 
Bengal, Mirpur, Dacca. 


288. Myxobolus sp. 

■fMyxobolus sp., Southwell, 1915, pp. 312-13, pi. xxvi, fig. 3. 
Myxobolus sp., Kudo, 1920, p. 151 ; 1933, p. 209. 

Cyst soft, flattened, roughly oval or lenticular, milky -white 
in colour. Spore with two equal polar capsules, with a very 
short anterior tail-Hke process. lodinophilous vacuole present. 

Dimensions. — Cysts up to 1-1 mm ; spores 13/x by 13/x ; 
polar capsule 4^ hy 4 [jl. 

Habitat. — Subcutaneous intermuscular tissue of Rasbora 
daniconius (Ham.-Buch.) from a stream near Katiwan : 
United Provinces, Mirzapore. 

Genus THELOHANELLUS Kudo, 1933. 

Myxobolus (part), Biitschli, 1882, pi. xxxviii ; Labbe, 1899, pp. 97- 
101 ; Kudo, 1920, p. 132 ; Wenyon, 1926, p. 727 ; Reichenow, 
1929, pp. 1068-72; Kudo, 1931, p. 312; Calkins, 1933, p. 568. 

Thelohanelhis, Kudo, 1933, pp. 198, 210. 

Spore pyriform ; flattened. One polar capsule at the 
anterior end. Sporoplasm with an lodinophilous vacuole. 
Histozoic in freshwater Fish. 

Key to Indian Species. 

1 (2). Cyst oval to cylindrical ; spore 30-32 /x [Prashad), p. 342. 

by 7-8 |u, T. rohitse (Southwell & 

2(1). Cyst elongated, ellipsoidal; spore [Prashad), p. 344. 

13-2-13-6jLt by 10-1-10-3/i T. seni (Southwell & 

289. Thelohanellus rohitae (Southwell & Prashad). (Fig. 169.) 

^Myxobolus rohitae, Southwell & Prashad, 1918, pp. 344-7, pi. xi, 
figs. 1-27. 
Thelohanellus rohitse. Kudo, 1920, p. 132 ; 1933, p. 210. 

Cysts oval to cylindrical, with the long axis of the cyst 
parallel to the gill -filaments, creamy-yellow in colour. The 
wall of the cyst is formed of a vertically striated portion 
showing no nuclei, covered externally by a membrane, two 
to three layers thick, and lined internally by an endoplasmic 
layer showing a coarse granular structure and with scattered 
nuclei. In the endoplasm two kinds of nuclei are found, 
viz., the vegetative and the generative. The latter occur 
in rounded cells called pansporoblasts or propagative cells. 
They are rounded in shape, with a marginally situated nucleus, 
and vary in size from 6 to 11 fi. The nucleus in the pan- 
sporoblast divides mitotically. Finally ten fully formed 
nuclei can be distinguished in the mother-pansporo blast, 
besides two nuclei for the pansporoblast mother-cell, and 



reduction nuclear chromatin particles lying free in the 
cytoplasm of the mother-cell. The pansporoblast ultimately 
divides into two daughter-cells or sporoblasts, each with five 
nuclei ; two of these unite later to form the nucleus of the 
sporoplasm, one forms the nucleus of the polar capsule and 
the other two are for the spore -membrane. A fully formed 
spore is an elongated pear-shaped body, rounded posteriorly 
and acutely pointed anteriorly. The spore-wall consists of 


Fig. 169. — Thelohanellus rohitse (Southwell & Prashad). A, gill with 
cysts ; B, young spore ; C, mature spore. (After Southwell 
and Prashad.) 

two valves ; the suture is a distinctly thickened ridge lying 
parallel to the long axis of the spore. There is only one 
polar capsule in each spore, with a long and much-coiled 
polar filament. An iodinophilous vacuole is present. 

Dimensions. — Cysts 3"l-3'8 mm. by 0'8-r2 mm. ; spores 
30-32 fjL by 7-8 ju, ; polar capsule 22-23 /x in length. 

Habitat. — Gills of Labeo rohita (Ham.-Buch.), from Turag 
Biver : Bbngal, Mirpur, Dacca. 


290. Thelohanellus seni (Southwell & Prashad). (Fig. 170.) 

■fMyxobolus seni, Southwell & Prashad, 1918, p. 347, pi. xi, figs. 28- 

Thelohanellus seni. Kudo, 1920, pp. 132-3 ; 1933, p. 210. 

Cyst elongated, ellipsoidal, whitish, with black scattered 
granules on the surface. Spore oval, pointed at the anterior 
end and much wider behind. Two valves, suture shghtly 
thickened. Polar capsule single, with a much-coiled filament, 
lodinophilous vacuole present. 

Fig. 170. — Spore of Thelohanellus seni (Southwell & Prashad). 
(After Southwell and Prashad.) 

Dimensions. — Cysts 4-7-5-4 mm. by 2-9-3-7 mm. ; spores 
13-2-13-6/i. by 10-l-10-3ft ; polar capsule about 4/x in length, 
filament 43/i in the extruded condition. 

Habitat. — On the median and caudal fins of Labeo rohita 
(Ham.-Buch.) : Bengal, Mirpur, Dacca. 

Genus HENNEGUYA Thelohan, 1892. 

Henneguya, Thelohan, 1892, pp. 167, 176 ; 1895, p. 352. 

Myxobolus (part), Gurley, 1893, p. 418 ; Cohn, 1896, pp. 227-72. 

Henneguya, Labbe, 1899, pp. 101-4 ; Minchin, 1903, pp. 286, 296; 
1912, pp. 409, 426 ; Kudo, 1920, pp. 158-73, 198-9 ; Wenyon, 
1926, p. 728; Reichenow, 1929, pp. 1059, 1072; Kudo, 1931, 
pp. 312-13 ; 1933, pp. 198, 210-12 ; Calkins, 1933, p. 568 ; 
Reichenow, 1935, p. 388. 

Spores ovoid al ; flattened. With a single or double caudal 
prolongation. Two pjTiform polar capsules at the anterior 
end. Monosporous, disporous or polysporous. Sporoplasm 
contains an lodinophilous vacuole. Mostly histozoic (a few 
coelozoic) parasites in freshwater Fish ; one species in 
a marine Fish. 

Key to Indian Species. 

1 (2). Cysts attached to the gill-filaments of [p. 345. 

the host H. ophiocephali Chak., 

2 (2). Cysts in the bulbus arteriosus of the [p. 345. 

host H. otolithi Ganapati, 


291. Henneguya ophiocephali Chakravarty. 

■fHenneguya sp., Ray, 1933 b, p. 349. 
\Henneguya ophiocephali, Chakravarty, 1938. 

Cysts spherical, attached to the gill-filaments of the infected 
host. Polysporous. Spores ovoidal or oblong, with the 
anterior end rounded and broader and the posterior 
tapering and forming a tail which is bifurcated along its 
entire length. Polar capsules elongate, with anterior end 
pointed and provided with a coiled filament, one capsule 
shghtly smaller than the other. An iodinophilous vacuole is 

Dimensions. — Cysts about 2 mm. in diameter ; spore 
16-4-20-5|Lt in length, 6-15 /x in breadth, tail 28-32 )[a ; polar 
capsules 6-18.U, by 2-06)u, and 5-18/u, by 2-06/Lt respectively; 
polar filaments 26-32/*. 

Remarks. — Description is based on Chakravarty's paper in 

Habitat. — Gills and muscles of Ophiocephalus punctatus 
Bloch : Bengal, Calcutta. 

292. Henneguya otolithi Ganapati. 

■fHenneguya sp., Ganapati, 1936, p. 204 ; 1938, p. 155. 

The affected area presents numerous white pustules, which 
are cysts containing the spores. Trophozoite shows a clear 
differentiation into ectoplasm and endoplasm, and vegetative 
and generative nuclei are present. Pansporoblasts originate 
by divisions of single generative cells. Each pansporoblast 
gives rise to two spores. Autogamy occurs. The phenomenon 
of dififuse infiltration seems to be much pronounced, bringing 
about considerable pathological changes. 

Remarks. — This is the second record of a Myxosporidian 
infecting the heart, the previous one being that by Keyssehtz 
of Myxoholus cordis from the ventricle of Barbus fluviatilis. 

Habitat. — Tissue parasite in the bulbus arteriosus of Otolithus 
ruber (Bl. Schn.) and 0. maculatus (Kuhl & Hass.) : Madras, 



The organisms included in this order are characterized by 
spores of comphcated structure. Each spore has its membrane 
or shell composed of three valves, which may be drawn out 
into simple or bifurcated processes. It has three polar 
capsules, and the polar filaments are visible in vivo. Several 
sporoplasms occur in each spore. 

They are parasitic in the body-cavity or the gut-epithelium 
of freshwater or salt-water Annelids, but have not been 
studied in India. 

Identification Table of Families. 

1 (2). Spore with a double membrane ; 

outer trivalve and the inner a 

single piece. A single binueleate [ = TetractinomyxidaB. 

sarcoplasm Haploactinomyxidse* Granata 

2 (1). Spore-membrane a single trivalve 

shell. A single eight-nucleate or [ = Triactinomyxidae. 

eight-uninucleate sporoplasm . . . EuactinomyxldaB* Granata 

Balbiani, 1882. 

Intracellular parasites typically of Arthropods and Fishes. 
Pseudopodia and amoeboid movements have rarely been 
observed. Multiphcation takes place by schizogony, and the 
resulting agametes are small, uninucleate bodies with indefinite 
outlines. Successive nuclear divisions, not accompanied 
by cell-division, lead to chain formation. These ultimately 
give rise to sporulating individuals or sporonts. Sporont 
develops into one to numerous spores. The spores are 
relatively small and less complex than those of Myxospobidia. 
They are ovoidal or bean-shaped. The spore-membrane is 
usually of a single piece and envelops the sporoplasm, and 
the polar filament is very long and fine. The filament may 
be contained in a polar capsuJe or lie coiled in the spore. 
In rare cases there may be two capsules and filaments in 
a spore. Owing to their small size, the polar capsules and 
filaments are invisible or obscure in the living spore, but 
can be rendered visible upon treatment with alkahes. Inter- 
mediate hosts are unknoA^Ti. 



The life-history of Stempellia magna, as described by Kudo, 
may be regarded as typical. When the spore reaches the 
mid -gut of the CuHcine host, the polar filament is extruded, 
the unicleate sporoplasm creeps out, enters a fat-cell, and 
reproduces by division (fig- 171, A). The products of this 
division become multinucleated, with four to eight nuclei (B) 
These chains then break up into binucleated cells, the nuclei 
of which discard some chromatin and fuse together in pairs (G) 
The sporonts thus formed may give rise to a single spore (D) 
or divide to form two (E), four (F) or eight (G) sporoblasts 

Fig. 171. — Diagram showing the probable development of a typical 
Microsporidian, Stempellia magna Kudo, X 800. A, amoe- 
bula coming out from the spore in the mid-gut of Culi- 
cine larva; B, stages in schizogony or nuclear increase ; 
C, sporont formation ; D-G, formation of one, two, four 
or eight sporoblasts ; H, transformation of a sporoblast 
into a nucleated spore with polar capsiile. (After Kudo . ) 

Each of these forms a single spore after chromidia formation 
and reconstitution of small nuclei has taken place (H). 

The Micro SPOEIDIA are responsible for causing devastating 
epidemics in silkworms, honey-bees, and certain fishes. AH 
the tissues of the host may become infected, and the tissue- 
cells destroyed on an extensive scale. Many species cause 
tumour-like masses to be formed in which the organisms may 



be encapsuled in a membrane derived from the host ; in others 
the membrane may be wanting. 

Perez (1905) subdivided the Microsporidia into two sub- 
orders, ScHizoGENEA (or Oligosporea) and Blastogenea (or 
Polysporea). In the former the principal trophic phase is 
a uninucleate meront which multiplies by fission, and finally 
gives rise to a sporont ; and in the latter there is a multi- 
nucleate Plasmodium producing sporonts by internal cleavage. 
As the exact structure of the trophic phase in many forms 

Fig. 172. — Effects of Microsporidian infection upon certain hosts. 
A, central nervous system of Lophius piscatorius infected 
by Nosema lophii (Doflein) ; B, fish showing a heavy 
infection by Glugea hertwigi Weissenberg ; C, a Culicine 
larva infected by Thelohania opacita Kudo, X 8 ; D, a 
Simulium larva infected by Thelohania multispora (Strick- 
land), X 6 ; E, F, normal and hypertrophied nuclei of 
the adipose tissue of larvse of Culex pipiens, the latter as 
afiectedhy Stempellia magna Jixido, X 750. (After Kudo.) 

is not known with certainty, the Microsporidia are now 
generally divided, following Leger and Hesse (1922), into 
two suborders, as follows :— 

1 . Spore with a single polar filament . 

2. Spore with two polar filaments ' . 

MonocnideaLeger&Hesse,p. 348. 
Dicnidea Leger & Hesse, p. 360. 

I. Suborder MONOCNIDEA Leger & Hesse, 

Microsporidia in which the spore is provided with one 
polar filament typically coiled in a polar capsule. 

Identification Table of Families. 

Spore oval, ovoid or pyriform ; if sub- 
cylindrical, length less than four times [p. 349. 

the breadth NosematidSB Labbe, 

Spore spherical or subspherieal Coccosporidse * Kudo. 

Spore tubular or cylindrical, length greater [Hesse, 

than five times the breadth Mrazekiidse* L^ger & 

NOSEMA. 349 

Family NOSEMATID^ Labbe, 1899. 

Spores oval, ovoid or pyriform ; if subcylindrical, length 
is less than four times the breadth. 

Key to Indian Genera. 

1 (2). Sporont becomes a single sporoblast [p. 349. 

forming a single spore Nosema Nageli, 

2(1). Sporont develops into eight sporoblasts [neguy, p. 356. 

and ultimately into eight spores Thblohania Hen- 

Genus NOSEMA Nageh, 1857, emend. Perez, 1905. 

Nosema, Nageli, 1857, p. 760. 

Glugea (part), Thelohan, 1895, p. 356. 

Myxocystis, Mrazek, 1897, pp. 1-5. 

Nosema (part), Labbe, 1899, pp. 105-8. 

Nosema, Minchin, 1903, p. 297 ; Perez, 1905, p. 17. 

Myxocystis (part), Mrazek, 1910, pp. 245-59. 

Nosema, Minchin, 1912, p. 418 ; Castellani & Chalmers, 1919, 
p. 529 ; Kudo, 1924 c, p. 65 ; Wenyon, 1926, p. 738 ; Knowles 
1928, pp. 324-30; Reichenow, 1929, p. 1088; Kudo, 1931, 
p. 320 ; Calkins, 1933, p. 569 ; Reichenow, 1935, p. 389. 

The vegetative form divides by multiple or binary fission 
into uninucleate rounded bodies or sporonts, each of which 
gives rise to a single ovoid or pyriform spore, which is not 
enclosed in a capsule. 

Key to Indian Species. 

Spores oval, 3 /x by 1-7 /n. In bed-bugs. N. adiei (Christophers), p. 349. 
Spores egg-shaped, 3-4 /x by 1-2 /x. In 

silkworms N. homhycis Nagel?, p. 351. 

Spores oval, up to 1-5 fi. In dog-fleas. N. ctenocephali Kudo, p. 354. 

293. Nosema adiei (Christophers). (Fig. 173.) 

fLeishmann-Donovan bodies, Adie, 1922 a, p. v. 
Bodies found by Mrs. Adie in salivary glands of bed-bug, Christo- 
phers, 1922, p. V. 
fBodies observed in Cimex rotundatus, Adie, 1922, pp. 236-8, 

pi. iv, figs. 1—4. 
^Nosemu adiei. Short & Swaminath, 1924, pp. 181-4, pi. xiii, figs. 

Nosemn adiei, Knowles, 1928, pp. 225, 330 ; Iyengar, 1929, p. 140. 

Spores elliptical or oval, with sharp and distinct boundary 
wall. They stain a rather pale, washed-out blue, with a central 
dot which may be of a deeper blue or red colour, and both 
kinds may occur in the same intracellular group. Planonts 
irregular in shape. Cytoplasm stains a bright shade of slate- 



blue, and possesses a large mass of chromatin, usually excentri- 
cally placed. Usually seen singly, and not definitely intra- 
cellular. Planonts come to rest in some suitable nidus, round 
off, forming meronts, which undergo schizogony. Meronts 
rounded or ovoid. Cytoplasm stains a cobalt blue and there 
may be one or two comparatively large masses of chromatin 
which may be central or excentric. 



Fig. 173. — Nosema adiei (Christophers). A, spores ; B, planonts ; 
C, meronts. (After Short and Swaminath.) 

Dimensions. — Planont about \'Q fx in diameter; meront 
3'2 /x by 2-7 ^ ; spore 3 |U by 1-7 ^. 

Habitat. — Gut, sahvary glands, and ovaries of Gimex 
rotundatus Linne : Assam. 

NOSEMA. 351 

294. Nosema bombycis Nageli. (Fig. 174.) 

Nosema bombycis, Nageli, 1857, p. 760. 

Panhistophyton ovatum, Lebert, 1858, pp. 149-86. 

Pebrine corpuscles, A. de Quatrefages, 1860, pp. 1-638 ; Pasteur, 
1870. pp. 1-327. 

Microsporidie, Balbiani, 1884, pp. 150-68. 

Pebrine corpuscles, Wood-Mason, 1887, pp. 1-3 ; Pfeiffer, 1888, 
pp. 469-86. 

Glugea bombycis, Thelohan, 1894, pp. 1425-7 ; 1895, pp. 357-8. 

Nosema bombycis, Labbe, 1899, pp. 106-7. 

Glugea bomJjycis, Toyama, 1900, pp. 1—40. 

Nosema bombycis, Minchin, 1903, p. 297 ; Leger & Hesse, 1907, 
p. 6 ; Stempell, 1909, pp. 281-358 ; Minchin, 1912, pp. 411, 
413-14, fig. 172; Omori, 1912, pp. 108-22; Kudo, 1913, 
pp. 368-71; 1916, pp. 31-51; 1918, pp. 141-7; 1924 c, 
pp. 69-76, figs. 1-39 ; fig. 757, text-figs. B2, D. 
^NosemM bombycis, Hutchinson, 1920, pp. 177-245, pis. i-xxvi. 

Nosema bombycis, Wenyon, 1926, p. 736, fig. 313 ; Knowles, 1928, 

pp. 324-30, figs. 72-5. 
'\Nosemxi bom,bycis, Iyengar, 1929, p. 140. 

Nosema bombycis, Reichenow, 1929, pp. 1088—90, figs. 1085—9 ; 
Kudo, 193i, p. 320, fig. 137, a, b ; Reichenow, 1935, p. 389, 
fig. 43. 

Spore egg-shaped, with the anterior end somewhat narrower 
than the other. It contains two vacuoles, one near the anterior 
and the other near the posterior end. The single polar capsule 
lies axially in the spore, occupying its whole length, and contains 
a long polar filament wound spirally in its interior. The sporo- 
plasm forms a cytoplasmic girdle round the polar capsule, 
separating the two vacuoles and placed slightly nearer the 
anterior pole of the spore. It contains at first a single nucleus 
which later divides into two and then into four. When the 
spores are ingested by a silkworm, the polar capsule ruptures 
in the lumen of its gut, and the polar filament is extruded, 
attaching the spore to an epithelial cell. This explosion of the 
polar capsule can be artificially brought about by treating the 
spores in some infected material with dilute acids or iodine 
solution, or even by pressure between coversHp and the shde. 
When germination of the spore takes place, the sporoplasm 
or amoebula creeps by amoeboid movement along the polar 
fiJament to an epithelial cell, which it invades. The planonts 
multiply by binary fission (fig. 174, A), become distributed 
throughout the body, enter tissue-cells, and become meronts (B). 
These meronts are spherical or oval in shape, and divide 
actively by fission (0) or by multiple division (D). The host- 
cell finally becomes completely filled with schizonts. Each 
meront ultimately develops into a spore {E). The spores are 
set free by the disintegration of the host-cells, and are taken 
with the food into the digestive tract of another host-larva, 
where the two nuclei of the spore divide once, giving rise to 
four nuclei (F). The polar filament is extruded (G) and later 



becomes detached. The binucleated sporoplasm creeps out 
through the opening of the capsule {H), and the remaining two 
nuclei degenerate in the spore. The two nuclei in the amoebula 
fuse into one and the uninucleate planont is formed. The 
entire life-cycle may be completed in four days. 

Dimensions. — Planont O-B-l-Sfiin diameter ; meront about 
3-5 ju. in diameter; spore 3-4 jj, by 1-2/x ; polar capsule 
1-5-2 /A by 0-8-1 /a; length of extruded filament 51-12 ^i, 
or even up to 98 ;u.. 

Pig. 174. — Life-cycle of Nosema bombycis Nageli. I, extracellular 
stages ; II, intracellular stages. A, planonts ; B, meront; 
C, binary fission of meronts; Z), multiple division of meronts ; 
E, stages in spore formation ; F, ripe spores in the mid- 
gut of a new host ; G, extrusion of the polar filament ; 
H, amcebula leaving the spore. (From Kudo, after 
Stempell. ) 

Remarks. — Ohshima (1927), discussing the function of the 
polar filament of the spore of Nosema hombycis, showed that 
as soon as filament extrusion is accomplished in the midgut 
of a silkworm the spore discharges a viscous fluid through 
the tube of the filament, which remains hanging in a spherical 
drop at the extremity of the filament. He has now shown 
(1937) that the polar filament is a germination tubule. In 
order to germinate the amoebula in the alimentary canal the 
spore evaginates its filament through the peritrophic membrane 

NOSEMA. 353 

into the epithelium, and the germ is safely poured out there 
through the tubule of the filament, protected on the way from 
the digestive ferment of the canal. The viscous fluid is the 
amcebula itself. The view previously held of regarding the 
polar filament as an attachment apparatus is thus con- 

Pathogenicity. — The organism causes a widespread disease, 
in silkworms, known as pebrine, or by various other names in 
different countries. In India it is locaUy known as Cota. 
The disease spreads among the host-larvse and to the offspring 
through the ova, causing disastrous epidemics and entailing 
huge economic loss to the silkworm industry. The outbreak 
of such disastrous epidemics in France and other countries 
of Europe, in the middle of the last century, led Pasteur to 
make his classical researches into the nature of the disease 
and the mode of its control. 

Lightly infected larvae do not show definite symptoms, but 
the heavily infected ones move about sluggishly, lose their 
appetite, grow slowly, and succumb to death before pupation. 
The cocoons spun by the infected larvee are very thin and 
poor in quaHty. In advanced stages of the disease irregular- 
shaped dark brown or black spots appear over the surface of 
the host-body, particularly on the posterior ventral side. 
The internal organs show a milky-white appearance due to 
the presence of a large number of spores. 

The eggs of the host may become infected while developing 
in the infected ovaries of the female moth, or the spores may 
be taken by the host-larvse into their digestive tract with 
contaminated food. The fsecal matter of the infected host- 
larvse is the most dangerous source of infection, as the con- 
tained spores are easily spread over the mulberry leaves on 
which the silkworms feed. The disease can, however, be 
controlled by eliminating both sources of infection. Infected 
eggs and all stages of the host-insect showing the least sign 
of infection must be destroyed as soon as possible by frequent 
and careful inspection. 

Hutchinson (1920) has made a study of the disease as it 
occurs in India. He has shown that pebrine spreads rapidly 
in Indian silk farms and is a much more acute problem 
than in Europe. He has further shown that light infections 
can be identified by dissecting out the gut of the moth and 
examining the posterior part of the colon, where fight infections 
are usually located. A far greater proportion of infected 
moths can thus be detected than by the Pasteur method. 

Habitat. — ^All tissues of eggs, larvse, pupse, and imagos of 
Bonibyx morii Fabricius, 

SPOK. 2 A 



295. Nosema ctenocephali Kudo. (Fig. 175.) 

■fNosenia pulicis, Korke, 1916, pp. 725-30, pi. Ixiii. 
Nosema ctenocephali, Kudo, 1924 c, pp. 105-6, pi. v, figs. 178-82 ; 
Iyengar, 1929, p. 140. 

Spore ova], refractile in fresh state, with one or two vacuolar 
spaces at the poles, and with an actively mobile sporoplasm. 


Fig. 175. — Stages in the life-cycle of Nosema ctenocephali Kudo. 

A, fully developed sporocyst (c.n., capsulogenous nucleus ; 
P.C., polar capsule ; p.f., polar filament ; p.n., parietal 
nucleus ; s., spore ; S.N., spore-nucleus ; s.t., sporocyst) ; 

B, germination with extruded filament, with a planont at 
its extremity (a, sporocyst ; b, planont ; c, same, highly 
magnified) ; C, colony of planonts ; D, division of meronts 
to form a chain-like group ; E, division of meronts 
to form a cluster ; F, capsule and polar filament (After 

NOSEMA. 355 

By addition of weak acetic acid or iodine water the movement 
within the spore is accentuated and results in the extrusion 
of the polar filament. The detached filaments are dehcate 
threads, sometimes nodular. The filament issues through 
the pore of the sporocyst and, as it breaks ofi", the amoebula 
emerges through the pore. In stained preparations the 
amoebulse are seen as minute bodies with irregular outhne, 
suggesting the possession of pseudopodial movement during 
life. The planont is a minute globular organism ; when 
active its movements are fairly rapid, a blunt pseudopodium 
being extruded in one direction only. Division takes place 
by binary fission. Planonts collect into groups or colonies 
of varying numbers of individuals. 

After penetration into a cell the mobile planont becomes the 
oval meront. The meront is larger than the previous phases, 
has homogeneous cytoplasm, with a centrally or excentricaUy 
placed refractile spot. It forms generations of merozoites, 
either by binary fission forming chain-like groups (fig. 175, D) 
or by multiple fission forming small clusters of cells (fig. 175, E). 
The final product of the division of the meront is a sporont 
which forms the sporocyst, polar capsule, and polar filament. 
The spore is recognized by its egg-shaped outHne and by the 
presence of one or two vacuoles at either pole. In stained 
films can be seen, in addition, a cystic wall containing two 
parietal nuclei, a polar capsule and polar filament, and the 
sporoplasm. The polar capsule is a hollow pear-shaped body 
containing a spirally twisted thread, and a capsulogenous 
nucleus situated at its broader end. The sporoplasm contains 
a, single nucleus and no vacuoles. 

Dimensions. — Planont 0-75-1 /a; spore up to 1-5/z, in length; 
length of extruded filament 25 ^u,. 

Remarks. — Heavily infected larvae are dark and mottled 
in appearance, owing to the presence of the Nosema, and are 
thus easily distinguished. 

Korke (1916) described the species as N. pulicis, probably 
in ignorance of the fact that NoUer (1912, 1914) had already 
described a species of that name from the dog-flea in Germany. 
As the dimensions of the spores are quite different in the 
two forms, Kudo (1924) has given the name iV. denocephah 
to the Indian form. 

Habitat. — Digestive tract of the larvae of the dog-flea, 
named by Korke Ctenocephalus felis (Bouche), probably 
Ctenocephalus canis (Curtis) : Punjab, KasauH. 




Genus THELOHANIA Henneguy, 1892. 

Thelohania, Henneguy, in Thelohan, 1892, p. 174; Gurley, 1893, 
p. 410; Thelohan, 1895, p. 361; Labbe, 1899, pp. 111-12; 
Minchin, 1903, pp. 285, 297, 320; Perez, 1905, p. 17 ; Minchin, 
1912, p. 418 ; Castellani & Chalmers, 1919, p. 529; Kudo, 1924 c, 
pp. 67, 130-63; Wenyon, 1926, p. 738; Reichenow, 1929, 
p. 1102 ; Kudo. 1931, p. 321 ; Calkins, 1933, p. 569 ; Reichenow, 
1935, p. 390. 

Each sporont develops into eight sporoblasts and ulti- 
mately into eight spores. The sporont-membrane may 
degenerate at different times of development. 

Key to Indian Species. 

1 (4). Spores oval, with equally rounded ex- 

tremities 2. 

2 (3). Spore 4-5-2 ju, by 2-4-2-8 fi T. indica Kudo, p. 356. 

3 (2). Spore 4-75-6 /x by 3-4 /n T. legeri Hesse, p. 357. 

4 (1). Spores broadly oblong with dissimilar 

extremities, showing a small knob at 
one end and a clear oblong space at 
the other ; 4-5-5 fj, by 3-3-5 ju T.ohscura Kudo, p. 359. 

296. Thelohania indica Kudo. (Fig. 176.) 

■^Thelohania sp., Iyengar, 1929, p. 138. 

\Thelohania indica. Kudo, 1929, pp. 3-4, figs. 19-50. 

The schizonts are elongated bodies containing four to eight 

Fig. 176. — Thelohania indica Kudo. A, tetranucleate schizont ; 
B, octonucleate schizont ; C, division into binueleate 
schizonts ; D-F, stages in formation of sporont ; G, spo- 
ront ; H, sporont with eight sporoblasts ; /, sporont with 
mature spores ; J, stained spore. (After Kudo.) 



nuclei arranged in pairs (fig. 116, A, B). They divide finally 
into two to four binucleate schizonts (C). The two nuclei 
become enlarged and closely associated, and ultimately fuse 
into one, thus forming a sporont (D-F). The sporont-nucleus 
undergoes three successive divisions, producing eight sporo- 
blasts. In each sporoblast the nucleus divides into two 
masses. One of them remains as a single nucleus of the 
sporoplasm, while the other disintegrates and produces the 
polar filament. The unstained spores are elhpsoidal, with 
equally rounded extremities ; the spore-membrane is moderately 
thick, and there is no indication of its being composed of two 
valves ; the contents are irregularly vacuolated. The stained 
spore shows a comparatively large nucleus at one end and 
a deeply stained polar capsule near the other. 

Dimensions. — Spores measure 4-5-2/^ by 2-4-2-8ju,. 

Habitat. — Adipose tissue cells of the larvae of Anopheles 
hyrcanus (Giles) : Bengal. 

297. Thelohania legeri Hesse. (Fig. 177.) 

Thelohania legeri, Hesse, 1904 a, pp. 570-1, 1904 6, pp. 571-2, 

10 text-figs. 
Thelohania illinoisensis. Kudo, 1921, pp. 167-71, 177 ; 1922, 

pp. 74-5. 
Thelohania legeri. Kudo, 1924a, pp. 147-62, 1 pi., 9 text-figs.; 

1924 c, pp. 143-6, figs. 499-507, 694-727, 764, text-fig. H ; 

Wenyon, 1926, p. 748. 
■\Thelohania sp., Iyengar, 1929, p. 138. 
\Thelohania legeri. Kudo, 1929, pp. 2-3, figs. 1-18. 
Thelohania legeri, Reichenow, 1929, pp. 1086, 1104; Kudo, 1931, 

p. 321, fig. 138, a-/; Calkins, 1933, p. 554, fig. 223. 

The earhest stages are unknown. The youngest stages 
found in the infected cells of the " fat-body " of the host are 
rounded bodies Avith compact chromatin granules (fig. 177, A). 
The nucleus becomes vesicular and then divides ; the cytoplasm 
becomes constricted and two uninucleated daughter-schizonts 
are formed {B). This division is repeated. Some of the 
binucleate schizonts do not spht, but grow into large elongate 
bodies with four nuclei (C). Each of these divides into two 
large binucleate schizonts. These nuclei lose their vesicular 
nature, become compact, and divide, the daughter halves 
remaining connected with each other by a strand. By further 
division two binucleate forms are formed, the nuclei in each 
being cousin nuclei and not daughter nuclei (D). Sometimes 
octonucleate schizonts occur, which by division produce 
four binucleate schizonts. The zygote or sporont is formed 
by the fusion of the two nuclei (E). The nucleus of the zygote 
divides three times in succession, resulting in stages with two 
(F), four (G), and eight nuclei (H). The sporont is then 
transformed into a pansporoblast with eight sporoblasts, 



each of which develops into a spore (/) . When the spore reaches 
the gut of a new host-larva the filament is extruded (J) and 
the sporoplasm emerges as an amoebula, starting the develop- 
ment again. 

Dimensions. — ^Meronts 3-4 ju, in diameter ; sporonts 9-1 O^a 
by 4-6 /x ; spores 4-75-6//. by 3-4/>(,. 

Remarks. — Iyengar (1929) described the life-cycle of Thelo- 
hania sp. from Anopheles pseudojam^esi Strickland & Chowd- 
hury {^=A. ramsayi Covell), and found it similar to T. legeri as 
described by Kudo. The measurements of the spores as 
recorded by him also fall within the limits of this species. 

Fig. 177. — Life-cycle of Thelohania legeri Hesse. A, youngest stage % 
B, formation of schizonts ; C, large schizont with four 
nuclei ; D, formation of binucleate forms ; E, sporont ; 
F, O, H, nuclear division of sporont to form binucleate, 
tetranucleate or octonucleate stages ; /, pansporoblast 
with eight sporoblasts ; ./, spore with extruded filament. 
(After Kudo.) 

Habitat. — Adipose tissue cells of larvae of Anopheles 
barbirostris Van der WuJp, A. annularis Van der Wulp {^A. 
fuliginosus Giles), A. varuna Iyengar, A. hyrcanus (Giles), 
A. ramsayi Covell, and A. subpictus Grassi {=A. rossi Giles) : 


298. Thelohania obscura Kudo. (Fig. 178.) 

■\Thelohania sp., Iyengar, 1929, p. 138. 
fThelohania obscura. Kudo, 1929, p. 4, figs. 51-61. 

Developmental stages not studied. Sporont octosporoblastic 
(fig. 178, il). Spores, when viewed in water after decolorization, 
are broadly oblong, often with dissimilar extremities, with 
a thin membrane containing within a rounded mass of homo- 
geneous and refractile protoplasm showing a small knob 
at one end and a clear oblong space at the other. The stained 

B C 

Fig. 178. — Thelohania obscura Kudo. A, sporont containing eight 
sporoblasts ; B, young spore ; C, mature spore. B and C 
stained with Giemsa's stain. (After Kudo.) 

spores show a nucleus near one end and a coiled polar filament 
forming an oblong mass {B, C). 

Dimensions. — Spores measure 4-5-5)u. by 3-3-5 /i. 

Remarks. — The record of T. legeri from India and the 
descriptions of T. indica and T. obscura are based on material 
sent to Kudo by M. 0. T. Iyengar from Bengal. As remarked 
by the latter, these parasites form whitish translucent masses 
beneath the cuticle of the thorax and the anterior abdominal 
segments of the larvae. The infected segments show hyper- 
trophy. None of the larvae showing these symptoms meta- 
morphosed, and all died before pupation. Those that are 
hghtly infected would be able to metamorphose into adults. 

Habitat. — Larva of Anopheles varuna Iyengar : BbngaIj. 

299. Gen. et sp. incert. 

Protozoan, Ross, 1906, p. 104. 

Thelohania (?) sp.. Kudo, 1921, p. 71 ; 1922, pp. 71-2. 

Gen. et spec, incert.. Kudo, 1924 c, p. 194. 

Eight spores were seen to be closely packed within an oval 
envelope. Spores were oval, refractive, apparently hard, with 
a circular vacuole at one focus of the elhpse. 

Habitat. — ^Nerve-cord of imago of Stegomyia sp. and Culex 
fatigans Wied. : India. 


II. Suborder DICNIDEA Leger & Hesse, 

Micro SPORIDIA in which the spore is provided with two 
polar capsules, one at each end, each containing a polar 

The suborder includes the single family Telomyxidse* 
Leger & Hesse, 1910, but no representative has been recorded 
from India up to the present time. 

Kudo, 1931. 

Minute spore, composed of a thin membrane of one piece, 
containing three uninucleate sporoplasms, around which is 
coiled a long, thick filament. Young trophozoites are histozoic 
or coelozoic, undergo schizogony, and produce uninucleate 
sporonts. The sporont divides twice and forms four small 
cells, each of which develops into a spore. 

The order has been created by Kudo (1931) to include the 
species Helicospiridium parasiticum Keihn from the body of 
certain insects. The structure of tbe spore is quite peculiar, 
and the species was previously referred to Haplospokidia. 
According to Kudo it seems to be best placed in the Cnido- 
SPORiDiA, if it is a Protozoan. 


Biitschli, 1882. 

These are typically parasites of the muscles of mammals, 
although birds and some reptiles are also known to harbom- 
them. They usually affect the striped muscles of the skeleton, 
tongue, larynx, and diaphragm. Lying between the muscle- 
fibres are thin- walled, whitish, tubular cysts termed Miescher's 
tubes, which are the remains of an infected muscle-fibre, 
the membrane of which is distended over the sac containing 
the parasites. They are generally visible to the naked eye, 
and often very large. When examined microscopically each 
is seen to consist of an envelope, from which partitions run 
inward, dividing the interior into a number of chambers 
containing vast numbers of crescent- shaped spores, the so-called 
" Raitiey's corpuscles." The spore is rounded at one end and 
pointed at the other ; it contains a nucleus near the rounded 
end and a collection of deeply staining chromatinic grains 
near the pointed end. Laveran and Mesnil described a striated 
structure at the pointed end which several workers considered 
as suggesting a rudimentary polar capsule, but recent authori- 
ties do not regard it as such. 

The spores are taken into the digestive tract of a specific 
host through the mouth. The spore -membrane ruptures, 
setting free the sporozoite, which enters the gut-epithelium. 
After multiplying there the organism makes its way into 
the muscular tissue. At first there is an elongate multi- 
nucleate mass, which may or may not divide into uninucleate 
bodies and which become the centre of infection in other 
muscle- fibres. Some of these trophozoites grow in size, and 
the body becomes divided into chambers, inside which spores 
are formed. 

Development has been studied chiefly in experimentally 
infected mice by Erdmann (1910, 1914), Negri (1910), and 
Crawley (1916), and in sheep by Alexeiefi" (1913). Smith 
(1901, 1905) was the iirst to demonstrate that mice could be 
infected by feeding them with the flesh of other infected 
mice. Negri (1910) and Darhng (1910 a) showed that guinea- 
pigs could be infected with the parasite of rats, and Darling 
also pointed out that the forms in guinea-pigs are morpho- 
logically identical with those from man. Erdmann (1910) 
infected mice with the parasite of sheep. Wenyon (1926) 


gives a list of thirty- nine species of Sarcocystis from a variety 
of hosts, but agrees with Alexeieff that there is no means 
of distinguishing the supposed species. 

Babudieri (1932) regards the Saecosporidia as more 
closely related to the Coccidia than the Cnidospoe.idia. He 
divides the subclass into Saecosporid-ia s. str. (including 
the famihes Sarcocystidse and Fibrocystidse) and Globidia, 
including the family Globididse. The Sarcocystidae includes 
the genus Sarcocystis with twenty-two species. The Fibro- 
cystidse includes Fihrocystis and Besnoitia with two species 
each. The Globididse includes Globidium with five species 
and Ileocystis with one species. The work should be referred 
to for the diagnostic features, synonymy, and hosts of various 
species. Krause and Goranoff (1933) tabulate the morpho- 
logical differences of forms described from birds. They 
succeeded in infecting the fowl with the strain from the 

Pathogenicity. — In many cases, even with fairly heavy in- 
fection, the host may not be adversely affected ; but in others 
the host sooner or later dies as the result of muscular degenera- 
tion. The organisms are known to produce a peculiar sub- 
stance called sarcocystin, which is highly toxic when injected 
into other animals. Teichmann and Braun (1911) showed that 
rabbits could be immunized against the toxin, and that the 
serum contained antibodies which could produce passive 
immunity in other animals. 

I. Order SARCOSPORIDIA s. s^r., Babudieri, 


Spore banana-shaped ; the cyst is wholly or partially of 
parasitic origin, and the spore produces metachromatic granules 
and secretes a toxin kno^oi as sarcocystin. The spore is 
produced by the prohferation of the pansporoblast, and the 
sporoblast has a peripheral situation in the cyst. 

Identification Table of Families. 

1 (2). Cysts elongated in form, parasitic in the [Babudieri, p. 363. 

striped muscle-fibres Sarcocystidae 

2 (1). Cysts rounded in form, parasitic in the [Babudieri. 

connective tissue or plain muscle-fibres . Fibrocystidse* 


Family SARCOCYSTID^ Babudieri, 1932. 
Genus SARCOCYSTIS Lankester, 1882. 

Sarcocystis, Lankester, 1882, p. 54. 

Sarcocystis -\- Balhiania-\- Miescheria, Blanchard, 1885, p. 244. 

Sarcocystis, Labbe, 1899, pp. 116-19; Minchin, 1903, p. 308 j 
Minchin, 1912, pp. 419-22 ; Castellani & Chalmers, 1919, 
pp. 530-3 ; Wenyon, 1926, pp. 760-9 ; Knowles, 1928, 
pp. 333-7 ; Reichenow, 1929, pp. 1120-7 ; Kudo, 1931, pp. 326-8 ; 
Calkins, 1933, pp. 555-6 ; Reichenow, 1935, p. 394. 

With the characters of the family. 

The cysts are to be found in practically all body -muscles ; 
in Ruminants they are found specially in the oesophagus, in 
other animals in the heart, intercostal or abdominal muscles. 
The cyst-wall may be produced entirely by the parasite 
(as in Sarcocystis muris) or in part by the host (as in S. tenella). 
The cyst is divided into chambers, the peripheral ones containing 
pansporoblasts or sporoblasts, while the central ones contain 
mature spores. The mature spores have neither polar capsules 
nor polar filaments. They show a very granular nucleus, 
a thin cell-membrane, and many metachromatic granules in the 

Many species have been recognized as occurring in man, 
sheep, cattle, horses, rats, and mice on the basis of occurrence 
in different host- species and shght difference in the dimensions 
of the spore ; but they are otherwise morphologically indis- 

300. Sarcocystis blanchardi Doflein, 1901. (Fig. 179.) 

Sarcocystis blanchardi, Doflein, 1901, p. 221, figs. 197, 198. 
"f Sarcocystis tenella, Shipley, 1904, pp. 45-7, pi. i, figs. 10, 17. 
'f Sarcocystis tenella bubalis, Willey, Chalmers, & Philip, 1904, 

pp. 65-72, figs. 1, 2. 
'ISarcocystis sp., Chatterjee, 1907, pp. 77-8. 
Sarcocystis tenella bubali, Wenyon, 1926, p. 768. 
Sarcocystis blanchardi, Wenyon, 1926, p. 768. 
Sarcocystis tenella, Wenyon, 1926, pp. 760, 768, fig. 329 ; Knowles,. 

1928, pp. 333-6. 
Sarcocystis blanchardi, Reichenow, 1929, p. 1123, figs. 1126, 1127 ;, 
1935, p. 392, fig. 46. 

Cysts somewhat pointed, though not very sharply, at either 
end, or both ends truncated, the thickest portion being in the 
middle. The covering of the cyst consists of two sheaths, the 
outer one being continuous and the inner forming partitions 
which divide the cyst into a number of polyhedral chambers 
with granular contents. The peripheral chambers are com- 
pletely filled with fine granulations and stain deeply. The 
central chambers are empty or contain spores. Spores are 
minute crescent-shaped bodies with granular contents. 

Dimensions. — The largest cysts measure 30 mm. in length,, 
5 mm. in breadth, and 3 mm. in thickness. 



Remarks. — Shipley (1904) and Willey, Chalmers, and 
Philip (1905) iirst described the sarcosporidial infection in 
buffaloes in Ceylon. According to Wenyon as many as four 
species have been named by different observers from the 
buffalo, but these are probably identical, and Reichenow 
recognizes only 8. hlanchardi. Chatter jee (1907) found the 
cysts and spores of Sarcocystis sp. in the heart-muscle of a cow. 
He did not find fine radiations round the capsule. The spores 

Fig. 179. — Sarcocystis hlanchardi Doflein. A, transverse section ; 
B, longitudinal section of the infected muscle-fibres of 
a young individual (a, muscle-fibre ; h, parasite) ; C, longi- 
tudinal section through a portion of the cyst in the oesopha- 
gus of cattle (a, muscle-fibre ; b, cyst-membrane ; c, nucleus 
of the muscle-fibre ; d, spores ; e, partition-walls between 
the chambers). (From Wasielewski, after Van Eecke.) 

were grouped in loculi, but distinct alveolar partitions could 
not be made out. 

Habitat.' — Muscles of buffalo, Bos huhalus Linn. : Ceylon ; 
heart-muscle of cow, Bos indicus Linn. : Bengal, Calcutta. 



301. Sarcocystis lindemanni (Rivolta). (Fig. 180.) 

?, Lindemann, 1863, p. 426 ; 1865, p. 382. 

Gregarina lindemanni, Rivolta, 1878, p. 12 ; Haden, 1883, p. 326 ; 

Blanchard, 1885, p. 244; Eve, 1889, p. 444; Target, 1889, 

p. 444. 
Sarcocystis hominis, Rosenburgh, 1892 ; Baraban & St. Remy, 

1894, pp. 79-82, figs. 1-5. 


Fig. 180. — Sarcocystis lindemanni (Rivolta). A, B, transverse sections 
of infected muscle-fibres ; C, individual spores. (After 

Sarcocystis lindemanni, Labbe, 1899, pp. 117-18 ; Minchin, 1903, 
p. 351. 
■[Sarcocystis sp.. Darling, 1919, p. 98. 
Sarcocystis lindemanni, Hegner & Taliaferro, 1924, pp. 373-4, 
figs. 139 h, 140 ; Wenyon, 1926, pp. 767-8. 


^Sarcocystis sp., Vasudevan, 1927, pp. 141-2, pis. xxiii, xxiv, 
figs. 1-9. 
Sarcocystis lindemanni, Knowles, 1928, p. 336, fig. 77, 2, 3; 
Reichenow, 1929, pp. 1123, 1125, fig. 1128; Kudo, 1931, 
p. 328 ; Reichenow, 1935, p. 394. 

The cysts (the so-called " Miescher's tubes ") are just 
visible to the naked eye as very thin, long, whitish streaks in 
the muscle-fibres and running parallel to them. The sheath 
proper is clear and homogeneous. External to this is a false 
sheath formed by compressed muscle-fibres showing clearly 
their nuclei and striation. From the true capsule numerous 
septa run into the cyst, dividing it into a number of partitions 
and giving a honeycombed appearance. On crushing one 
of these cysts numerous tiny curved bodies or spores are 
liberated. These spores are somewhat crescentic and pointed 
at both ends. No terminal filaments could be seen. The 
nucleus in each spore is oval, placed along the long axis 
either in the centre or, more commonly, subterminally. In 
some spores near one end is a stained area which at one time 
was supposed to represent a polar capsule. No vacuole is 

Dimensions. — In the case of human sarcosporidiosis observed 
by Baraban and St. Remy Miescher's tubes were from 150 to 
1600/x long and from 77 to 168|L(, in thickness ; Vasudevan 
gives 5-3 cm. as length and 322 ^u, as thickness. Spores are 
8-33/x in length and 1-66 ^u. in width, the nucleus in the spore 
measuring 3-33//, by 1-66/x,. 

Remarks. — The cysts of this parasite are much thinner and 
longer than those of S. miesheriana, and are just visible to the 
naked eye. The spores also are smaller and both ends are 
pointed instead of one end being blunt as in that species. 

The parasite described by Vasudevan (1927) is of special 
interest, as only six or seven cases of sarcosporidial infection 
have previously been recorded in man. 

Habitat. — ^Muscles of a man : Madbas, Madras. 


II. Order GLOBIDIA Babudieri, 1932. 

Spores fusiform containing large siderophil granules in the 
centre. Cysts occur exclusively in the intestinal submucosa, 
inside a hypertrophied host-cell, with a peripheral nucleus. 
No toxin secreted. 

Family GLOBIDID^ Babudieri, 1932. 

With the characters of the order. 

Genus GLOBIDIUM Flesch, 1884. 

GloUdium, Flesch, 1884, p. 459; Labbe, 1899, p. 72; Minchin, 

1903, p. 350. 
Gastrocystis, Chatton, 1910, p. 114. 
Besnoitia, Franco & Borges, 1916, p. 269. 
Globidium, Wenyon, 1926, pp. 769-73 ; Knowles, 1928, pp. 337-8 ; 

Reichenow, 1929, pp. 905-7 ; 1935, p. 394. 

Cysts spherical, up to 5 mm. in diameter, embedded in the 
mucosa of the aUmentary canal or skin of mammals. Each is 
enclosed by a membranous capsule and, when full grown, con- 
tains groups of spores which resemble those of Sarcosporidia. 

Remarks. — Flesch (1883) observed the infection in the horse ; 
Chatton (1910) in sheep and goats ; Gilruth and Bull (1912) 
in the kangaroo, wallaby, and wombat ; Kupke (1923) in the 
horse ; Wenyon and Scott (1925) in the wallaby ; and Henry 
,and Masson (1932) in the camel. There is great divergence 
of opinion as regards the systematic position of the genus. 
Wenyon (1926) considers it as closely related to Sarcosporidia, 
Reichenow (1929) places it in an appendix after Gregarinida, 
and Henry and Masson (1932) regard the genus as large Coc- 
€IDIA, while Hobmaier (1922) beUeved that the parasites 
included in the genus are fungi, and not Protozoa at all. 
Babudieri (1932) placed it in a separate order of the subclass 
Sarcosporidia. Enigk (1934) has described the development 
of macrogametes, microgametes, and schizonts of Globidium 
■cameli and pointed out the similarity to that of Coccidia. 

302. Globidium fusiformis Hassan, 1935. (Fig. 181.) 

■\Globidiuni fusifortnis, Hassan, 1935, pp. 1-7, pis. viii-x ; Ware, 
1937, p. 35.' 

Spherical protoplasmic bodies, varjdng in size, the smallest 
with a single nucleus and the largest with as many as forty- 
eight or more nuclei in a single optical plane. The parasite is 
at first intracellular in the gastro-intestinal epithehal cells, 
but later the tissues form a thin capsule round the mass. 
The colony becomes intracellular and is pushed towards the 
subepithelial connective tissue. Ultimately the cysts are 



liberated in the lumen of the organ. The centrally situated 
single nucleus grows and undergoes division, and the many 
nuclei become arranged round the periphery of the cyst. 
Finally projections representing the extremities of marginally 
situated spores appear to radiate round the periphery of the 
spherical body. In a freshly ruptured cyst twenty or more 
spores are seen lying parallel to one another. Spores are 
elongate, spiadle-shaped, and sHghtly curved, one end being 
more finely pointed than the other. The nucleus is oval, 
vesicular, and situated near the rounded end of the spore. 
In the centre of the spore is a round chromatin mass which 
stains intensely, and minute granules of chromatin are also 
irregularly distributed on both sides of the nucleus. 

Fig. 1 81. — GlobidiumfusiformisHsiSsan. A, multinucleated cytoplasmic 
bodies ; B, a stage in the development of the spores ; 
C, cluster of spores from a freshly ruptured cyst ; D, in-- 
dividual spores. (After Hassan.) 

Dimensions. — Cysts measure 60-90 /x in diameter ; spores 
are 13/Lt by 2-2-5 fi, the nucleus measuring 3-25 ^u, by 2-25 //,. 

Remarks. — ^The parasite was examined in the dried faeces of 
a cow, and it is not certain if the form is morphologically 
distinct from G. besnoiti of the cattle and G. faurei {G. gilruthi) 
of the sheep and goats. 

Habitat. — ^Faeces of cow. Bos indicus Linn. : Punjab, Gurgaon. 

303. Globidium sp. 

Sarcocystis macropodis, Gilruth& Bull, 1912, pp. 432-50, pis. Ixxii- 

Globidium sp., Wenyon, 1926, p. 773. 
f Globidium sp., Knowles, 1928, p. 337. 

Remarks. — Gilruth and Bull (1912) recorded the infection 
in the kangaroo, wallaby, and wombat in Austraha. Knowles 
(1928) mentions that Acton observed a case of infection in 
a wallaby in the Zoological Gardens, Alipore, Calcutta, but 
a description does not appear to have been pubhshed. 

Habitat. — Alimentary canal of a wallaby : Bengal, Calcutta. 


IV. Subclass HAPLOSPORIDIA Liihe, 

Haplosporidia are characterized by the presence of large 
spores, each containing a single voluminous nucleus, but no 
polar capsule, and showing a simple type of development. The 
young parasite is an amoebula, which at first multipHes by 
fission. The nucleus of each daughter-cell multiplies repeatedly 
and produces a multinucleate plasmodium. The plasmodium 
may divide (plasmotomy) or produce merozoites (schizogony) 
or form spores. The spores are produced from sporoblasts, 
each forming a single spore, or from pansporoblasts, each 
giving rise to a number of spores. Infection is carried by the 
spores, which produce amoebulse and start a new generation. 

The spore is spherical or elhpsoidal, and is covered by 
a resistant membrane which may be marked by ridges or 
tubercles, and may be prolonged into a tail-hJke process. 
In a few species the spore-membrane possesses a hd, which 
opens to allow the uninucleate sporoplasm to emerge as an 

The subclass includes parasites of certain Invertebrates, 
chiefly AnneHds, and Fish. They float freely in the body- 
cavity fluid of the Invertebrates or infest the tissue- cells, such 
as those of the intestine. In Fish they attack the gills or other 
tissues, giving rise to white nodules. One species occurs 
in the Malpighian tubules of the cockroach. The genus Ehino- 
sporidium, species of which form httle cysts in the nasal 
cavities of man and horse, used also to be included in this 
subclass, but the studies of Ash worth (1923) have shown that 
it is not a Protozoon but a fungus. 

Debaisieux (1916, 1920) has shown that certain species of 
Haplosporidium CauUery & Mesnil and Bertramia Caullery & 
Mesnil should be included with the Microspokidia, while 
Coelosporidium Crawley is not a Sporozoon. In his view 
Haplosporidia should be abandoned as a separate group. 

For classification of the Haplosporidia, Caullery and 
Mesnil (1905) and Ridewood and Fantham (1907) may be 
referred to. The latter divided the Haplosporidia into two 
sections — ^the Oligosporijlea, in which each pansporoblast 
produces only a small number of spores or a single spore, 
and the Polysporulea, in which the pansporoblast gives rise 
to many spores, either successively or almost simultaneously. 
No one has as yet studied Haplosporidia in India. 

SPOR. 2 B 


Doubtful Protozoa. 

Genus RHINOSPORIDIUM Minchin & Fantham, 1905. 

Rhinosporidium, Minchin & Fantham, 1905, pp. 521-32 ; Minchin, 
1912, pp. 424-5 ; Castellani & Chalmers, 1919, pp. 533-4 ; 
Ashworth, 1923, pp. 301-42 ; Hegner & Taliaferro, 1924, pp. 375- 
7 ; Wenyon, 1926, pp. 776-8 ; Knowles, 1928, pp. 338-42 ; 
Reichenow, 1929, pp. 1136-7. 

Organisms giving rise to polypi, especially in the nose, of 
human beings and horses. Believed for a long time to belong 
to Haplosporidia, but now generally recognized as a fungus. 

304. Rhinosporidium seeberi (Wernicke). (Fig. 182.) 

Coccidium seeberi, Wernicke, 1900. 

?, O'Kinealy, 1903 a, pp. 109-12 ; 1903 b, pp. 43-4, 1 pi. 
■\Rhinosporidiuni kinealyi, Minchin. & Fantham, 1905, pp. 521-32, 

pis. XXX, xxxi. 
Rhinosporidium seeberi, Minchin, 1912, pp. 424—5. 
■f Rhinosporidium kinealyi, Tirumurti, 1914, pp. 703-19. 
■f Rhinosporidium seeberi, Castellani & Chalmers, 1919, pp. 533-4, 

fig. 190 ; Ashworth, 1923, pp. 301-42, 5 pis. 
Rhinosporidium seeberi, Hegner & Taliaferro, 1924, pp. 375—7, 
fig. 141 ; Wenyon, 1926, pp. 777-8, figs. 335, 336 ; Reichenow, 
1929, pp. 1136-7, fig. 1141. 

The youngest forms are spherical bodies, about 6 ju in diameter, 
embedded in the cytoplasm of connective tissue cells, with 
chitinoid envelope, vacuolated cytoplasm, and a vesicular 
nucleus containing a karyosome. Growth is accompanied 
by nuclear multiphcation and the cytoplasm becomes laden 
with numerous food-granules. When the parasite is about 
100/x in diameter and has about 128 nuclei the chitinous en- 
velope becomes much thickened by the deposition of cellulose 
on the inner surface, except at one point where the future 
pore will be formed. Nuclear divisions contitiue, and when 
there are about 4,000 nuclei the cytoplasm divides into rounded 
cells, which divide twice to form about 16,000 young spores 
within the sporangium. In the mature sporangium there are 
usually fully developed spores in the centre, and small, immature 
ones arranged peripherally. The fully-formed spore has a 
chitinous envelope, a vesicular nucleus with a karyosome, 
and cytoplasm containing one to sixteen refringent spherules 
of reserve food material. The spores are spherical or oval. 
They are discharged in hundreds through the pore of the 
sporangium, and are scattered throughout the connective 
tissue by the lymph exudate. The reserve food material is 
gradually used up and the spores enter into fresh connective 
tissue cells to repeat the cycle. The ripe sporangia are seen 
as white dots covering the surface of the nasal polypus. 



Dimensions. — Ripe sporangia 250-300 /a in diameter ; spores 
7-1 OjU. in diameter. 

Remarks. — The organism has been found in polypoid growths 
in nose, naso-pharynx, uvula, conjunctiva, lacrymal sac, 
ear, and penis of man. 

Tirumurti (1914) recorded fifteen cases from man, aflfecting 
various parts of the body, in the Madras Presidency. 


Fig. 1S2. — Rhinosporidium seeberi (Wernicke). A, section of stage 
with about 500 nuclei, showing the outer chitinous and 
inner cellulose layer of the envelope and the position 
of the future pore ; B, section of the stage in which the 
contents of the sporangium have divided into about 
4,000 nucleated cells ; C, discharge of mature spores ; 
D, section of spore showing nucleus with karyosome, 
and vacuolated cytoplasm. (After Ashworth.) 



Researches of Ashworth (1923) apparently prove that the 
organism belongs to the fungi, and ought to be classed among 
the Phy corny cetes. 

Habitat. — Polypi in the nose of man : Bengal, Calcutta ; 
Madras, Madras ; Ceylon. 

305. Rhinosporidium sp. 

flSTot identified, Vasudevan, 1932, pp. 299-302, pi. viii. 

Young trophozoite a spherical encapsuled body, with 
a distinct nucleus and nucleolus. The capsule is hyaline, 
doubly-contoured, and l-6ju, in thickness. The cjiioplasm is 
vacuolated and in some trophozoites contains thin protoplasmic 
granules. Nucleus central, occasionally peripheral and mar- 
ginal, 4'8jLt in diameter, with nucleolus 1-6 /x in diameter. In 
later stages the organism is larger and protoplasmic strands and 
granules are pronounced. One of them was somewhat pear- 
shaped, probably due to an obhque section. 

Dimensions. — The organism measured 40 ju, by 23^. 

Remarks. — ^The organism somewhat resembles the earlier 
stages ofi?^.mospon(Zmm in shape, hyaline capsule, protoplasmic 
strands, and granules, but differs from Rhinosporidium seeberi 
in that the earhest stage is much larger. The later stages, with 
spore-bearing sporangia, were not found. The infection was 
in the skin and not in a mucous membrane. 

Habitat. — Inside walls of abscess cavities, and sometimes 
in the chronic inflammatory fibrous tissue in man : Ceylon. 

306. Rhinosporidium equi Zschokke. 

Rhinosporidium equi, Zschokke, 1913, pp. 641-50 ; Hartmann, 
1921, p. 1387 ; Wenyon, 1926, p. 778. 
'\ Rhinosporidium sp., Rao, 1938, pp. 263-4. 
\ Rhinosporidium equi, Sahai, 1938, p. 264. 

Probably identical with R. seeberi from man. 

Remarks. — According to Sahai (1938) the first case in an 
equine was recorded by Krishnamurti Ayyar in Madras in 
1932. Sahai has recorded another case in a comitry-bred 
mare in Orissa. The animal had noisy breathing and a blood- 
stained mucous discharge oozing from one of the nostrils. 
There was a small cauliflower-Hke growth about one inch 
long by half an inch thick situated in the anterior part of 
the nasal chamber, shghtly obstructing the passage. In 
the connective tissue were found numerous cysts or sporangia 
in various stages of development, the fully matiu-e ones biu-sting 
to discharge the spores. Wenyon (1926) doubted if the 
equine form is distinct from the human form, and Sahai (1938) 
supports his view. 

Habitat. — Nasal cavities of Equus caballus Linn. : Madras, 
Madras ; Orissa, Bargarh (Sambalpur). 


307. Rhinosporidium sp. 

■fRhinosporidium sp., Rao, 1938, p. 263. 

Probably identical with R. seeberi from man. 

Remarks. — According to Rao (1938), Rhinosporidiosis was 
recorded a few years ago affecting two buUocks, one cow, 
and one pony in Madras, and has since been diagnosed in 
eighteen buUocks and one pony in the same province. Rhino- 
sporidiosis in bovines appears so far to have been reported 
from the Madras Presidency only. The lesions are found 
in the nose, and the presence of a trauma in the nose appears 
to be necessary for the development of the lesions. 

Rao thinks that the causal organism may be the same 
as in man, and suggests that the infection to man and animals 
may be through the dust raised while ploughing fields manured 
with bovine dung, as a large number of cases are met with in 
man and cattle engaged in agriculture. 

Habitat. — Nasal cavities of Bos indicus Linn. : Madras 




(i) Supplementary List of Parasites and their Hosts. 



Fam. Stomatophoeid^ (p. 

Stomatophora primitiva . . 

Fam. Lectjdinid^ (p. 18). 

Lecudina eunicse 

Leeudina lysidicse 

Fam. Gregakinid^ (p. 19). 

Ulivina eunicse 

Deuteromera cleava .... 
Contortiocorpa prashadi . . 

Fam. Sei-enidiid^ (p. 19). 
Selenidium amphinomi . . 


Fam. EiMEKiiD^ (p. 21). 
Eimeria arloingi 



Eimeria cylindrica 
Eimeria gupti . . . . 
Eimeria hoormse . . 
Eimeria labbeana . . 
Eimeria smithi . . . 

Eimeria wassilewskyi 
Eimeria zlXrnii 

Incert^ sedis (p. 22). 
Taxoplasma butasturis . . . 


Fam. H^MOPROTiED^ (pp. 
Hcemoproteus (?) halcyonis 

Hsetnoproteus (?) lanii . . . 

Haemoproteus ( ? ) therei- 

cerycis var. zeylonica . . . 
Leucocytozoon ardeolee . . . 
Leucocytozoon enriquesi . . 
Leucocytozoon molpastis . . 
Leucocytozoon (?) sp. ... 
Leucocytozoon (?) sp 


Eunice siciliensis .... 


Eunice siciliensis .... 


Lysidice collaris 


Eunice siciliensis .... 


Eunice siciliensis .... 


Eunice siciliensis .... 


Amphinome rostrata . . 

Coelomic cavity. 

Capra hircus 

Alimentary canal, 
Alimentary canal. 
Alimentary canal. 

Ovis sp 

Bos indicus 

Natrix piscator 

Lissemys punctata .... 
Columba livia 

Bos bubalus 

Bos indicus 

Axis axis 

Bos bubalus 

Bos indicus 

Alimentary canal 
Alimentary canal. 
Alimentary canal. 
Alimentary canal. 
Alimentary canal. 

Butastur teesa 



Halcyon smyrnensis 



Lanius schach erythro- 



Thereiceryx viridis. . . . 


Ardeola grayii 

Chloropsis jerdoni .... 
Molpastes cafer cafer . . 
Oriolus oriolus kundoo. 


Oriolus xanthornus xan- 




Parasite. Host. Seat. 

Fam. Plasmodiid^ (pp. 24-5). 

Proteosoma centropi Centropiis sinensis par- Blood. 

Proteosoma heroni Pond heron Blood. 

Proteosoma passeritse .... Passerita mycterizans 

Laverania malarias Anopheles varuna . . . 

Plasmodium, malarise .... Anopheles varuna . . . 

Plasmodium vivax Anopheles varuna . . . 


Fam. Theileeud^ (p. 25). 

Theileria hirci Ovis sp Blood. 

Theileria sp Bos indicu^s Blood. 

Fam. BABEsnD^ (p. 25). 

Babesia bovis Bos indicus Blood. 

Babesia taylori Capra hircus Blood. 

Inceet^ sedis (p. 26). 

Anaplasma marginale .... Bos indicus Blood. 

Bertarellia calotis . Calotes versicolor major. Blood. 

Bertarellia sp Lissemys punctata Blood. 



Fam. Ceeatomyxid^ (p. 26). 

Ceratomyxa hilsse Hilsa ilisha Gall-bladder. 

Fam. Myxidiidje (p. 26). 

Myxidium glossogobii . . . Glossogobius giuris .... Gall-bladder. 

Fam. Myxobolid^ (p. 26). 

Myxobolus calbasui Labeo calbasu Gall-bladder. 

Labeo rohita Gall-bladder. 

Myxobolus mrigalse Cirrhina mrigala Scales. 

Incert^ sedis (p. 27). 

Rhinosporidium equi . . . JEquus caballu^s Nasal cavities. 

Rhinosporidium sp Bos indicus Nasal cavities. 

(ii) Supplementary List of Hosts and their Parasites. 

Host. Parasite. Seat. 

Mammalia (pp. 27-9). 

Axis axis Eimeria wassilewskyi . Alimentary canal. 

Bos bubalus Eimeria smithi Alimentary canal. 

Eimeria zilrnii Alimentary canal. 

Bos indicus Eimeria cylindrica . . . Alimentary canal. 

Eimeria smdthi Alimentary canal. 

Eimeria ziirnii Alimentary canal. 

Theleria sp Blood. 

Babesia bovis Blood. 

Anaplasma tnarginale . Blood. 

Rhinosporidium sp. . . Nasal cavities. 

Bosefhalus tragocamelus . Eimeria yakomovi .... Blood. 

Capra hircus Eimeria arloingi Alimentary canal. 

Babesia taylori Blood. 

Equus cabalus Rhinosporidium equi . Nasal cavities. 

Ovis sp Theileria hirci Blood. 



AvES (pp. 29-31). 

Ardeola grayii 

Butastur teesa 

Centropus sinensis parroti 

Chloropsis jerdoni 

Columba livia 

Halcyon smyrnensis jusca . 

Lanius schach erythronotus 
Molpastes cafer cafer 

Oriolus oriolics kundoo . . . 
Oriolus xanthornus xan- 

Pond heron 

Thereiceryx viridis 

Reptilia (pp. 32-3). 
C'alotes versicolor major . . . 
Lissemys punctata granosa 

Natrix piscator 

Passerita tnycterizans .... 

Pisces (p. 34). 

Cirrhina mrigala 

Glossogobius giuris 

Hilsa ilisha 

Labeo calbasu 

Labeo rohita 

Otolithus maculatus 



Insecta (p. 35), 
Anopheles varuna . . . 


POLYCH^TA (p. 36). 

Amphinoine rostrata 
Eunice siciliensis .... 

Leucocytozoon ardeolse . 


Toxoplasma butastvris . 


Proteosoma centropi . . 


Leucocytozoon enriquesi 


Eimeria labbeana .... 


Hxmoproteus (?) hal- 


cyonis fvscm. 

Hsemoproteus (?) lanii . 


Leucocytozoon molpas- 

Leucocytozoon (?) sp. . . 



Leucocytozoon (?) sp. . . 


Proteosoma heroni .... 


Hsemoproteus (?) therei- 



Bertarellia calotis .... 


Bertarellia sp 


Eimeria gupti 


Protesoma passeritse . . 


Myxobolus mrigalse . . 


Myxidium, glossogobii . 


Ceratomyxa hilsse .... 


Myxobolus calbasui . . . 


Myxobolus calbasui . . . 


Henneguya otolithus . . . 

Bulbus arteriosus 

Laverania malaria . . . 


Plasmodium malarise . 


Plasmodium vivax . . . 


Lysidice collaris 

Selenidium amphinomi. Ccelomic cavity. 

Stomatophora primitiva Intestine. 

Lecudina eunicse Intestine. 

Ulivina eunicse Intestine. 

Deuteromera cleava . . Intestine. 

Contortiocarpa pra- Intestine. 


Lecudina lysidicse .... Intestine. 



Genus STOMATOPHORA Drzewecki, 1907, emend. 
Hesse, 1909, and Bhatia, 1924. 

(Pages 68-72.) 
308. Stomatophora primitiva Bhatia & Setna. (Fig. 183.) 

fStoinatophora sp., Bhatia & Setna, 1935, p. 312. 

J Stomatophora primitiva, Bhatia & Setna, 1938 (in press). 

Trophozoite almost spherical, Avith a cup-like sucker near 
the anterior end. The sucker is not provided with a central 

Fig. 183. — -Stomatophora primitiva Bh. & Set. 
(After Bhatia and Setna.) 

mucron. Numerous epicytal striations radiate outwards 
over the body for some distance round the sucker. Nucleus 
rounded, with a large spherical karyosome. Gametocysts 
slightly ellipsoidal. 

Dimensions. — Length of trophozoite 222 ju., width 203*5 ju, ; 
nucleus 44-4 /a by 37/x, : gametocyst measures 222 /x by 203-5 /i. 

Remarks. — ^All the previously known species of the genus 
Stomatophora and of other genera included in the family 
Stomatophoridse are parasites of earthworms belonging to 
the genus Pheretima. The present species is the first record 
from a polychsete. It recalls in its structure S. simplex Bhatia, 
1924, from which it differs in the absence of a central mucron 


in the sucker. Seen under the oil-immersion lens the sucker 
is seen to be surrounded by a clear area, and the surrounding 
epicytal striations appear to run into each other and interlace. 

The sucker was seen to be retained in the case of one of the 
gametocytes in a gametocyst. 

Habitat. — ^Intestine of Eunice siciliensis Grube : Andaman" 
Islands, Port Blair. 

Genus LECUDINA Mingazzini, 1891. 

(Pages 83-5.) 
Key to Indian Species. 

1 (3). Cytoplasm of the anterior portion of the 

body differentiated 2. 

2. Trophozoite cylindrical ; epimerite 

variable in form, with an anchor plate [Aiyar, p. 83. 

at the end L. brasili Ganap. & 

3(1). Cytoplasm of the anterior portion of the 

body not differentiated 4. 

4 (5). Trophozoite elongate oval ; epimerite [p. 378. 

knob-like L. eunicse Bh. & Set., 

5 (4). Trophozoite oval ; epimerite a long [p. 379. 

conical trunk L. lysidicse Bh. & Set., 

309. Lecudina eunicse Bhatia & Setna. (Fig. 184.) 

■fLecudina eunicse, Bhatia & Setna, 1938 {in press). 

Trophozoite elongate oval, widest in the middle, narrowing 
at both ends, bluntly pointed anteriorly, and more sharply 

Fig. 184. — Lecudina eunicse Bh. & Set. 
(After Bhatia and Setna.) 

pointed posteriorly. Epimerite forming a knob-hke structure, 
with its cytoplasm differentiated from that of the rest of the 
body. Nucleus large, subspherical, with a single karyosome. 

Dimensions.- — Length of the trophozoite 475 /x, maximum 
width 94- 5 /x ; epimerite 14ju, in length ; nucleus 38-5 ju, by 
28 /x. 

Remarks. — The parasite somewhat resembles L. elongata 
(Mingazzini) in the differentiation of its anterior end, which 
is described by Mingazzini as a small spherical button, but 
according to Reichenow (1932) an epimerite is absent, and the 


parasite attaches itself to the epithelial cells of the host 
by its differentiated anterior portion in a sucker-like manner. 
L. eunicss differs from L. elongata in the form of the tropho- 
zoite and the structure of its nucleus. 

Habitat. — Intestine of Eunice siciliensis Grube : Andaman 
Islands, Port Blair. 

310. Lecudina lysidiese Bhatia & Setna. (Fig. 185.) 

■\Lecudina lysidiese, Bhatia & Setna, 1938 (in press). 

Trophozoites oval, broadly rounded anteriorly and narrower 
and pounded off posteriorly, broadest in the anterior third 
of its body. Cytoplasm does not show any differentiation 
in the anterior portion of the body. Epimerite in the form 
of a long conical trunk. Nucleus spherical, oval or irregularly 
quadrilateral, with a large central karyosome. Sporonts much 
larger in size, without an epimerite, but showing a triangular 
area slightly raised from the anterior end of the body. 

Fig. 185. — Lecudina lysidiese Bh. & Set. 
(After Bhatia and Setna.) 

Dimensions. — Cephalonts with a total length of 71*9- 
185/x., maximum width 14- 8-62-9 /x, length of epimerite 11-1- 
37 /x, and nucleus 14-8 /i by 10 /x. Sporonts may reach a length 
of 370 /x, with a maximum width of 155-4 jix. 

Remarks. — The parasite invites comparison with L. aphroditse 
(Lank.), but differs in its much smaller size, in the nuclear 
structure, and also in the epimerite not being marked with 
successive constrictions. The nucleus varies considerably 
in form and position. It may be spherical, oval or irregularly 
quadrilateral in form, and may be situated in the anterior 
middle or posterior part of the body, but it always presents 
the same structure. The central karyosome is surrounded 
by a clear area, and chromatin particles are arranged in 
a peripheral zone within the nuclear membrane. The sporonts 
frequently and the cephalonts occasionally are seen to be 
full of elongated spindle-shaped bodies, each containing 
a definite central nucleus ; these are probably algse, and 
measure up to 74 /x by 3-7 /x. 

Habitat. — Intestine of Lysidice collaris Grube : Andaman 
Islands, Port Blair. 


Family GREGARINID^ Labbe, 1899. 

(Pages 96-107.) 

Genus ULIVINA Mingazzini, 1891. 

Vlivina, Mingazzini, 1891, p. 235 ; Labbe, 1899, p. 34 ; Minchin, 

1903, pp. 203, 325. 
Doliocystis (part), Crawley, 1903, p. 56. 
Ulivina, Saint-Joseph, 1907, pp. 164, 174. 
Doliocijstis (part), Ellis, 1913, p. 287. 

Ulivina, Kamm, 1922, pp. 28-9 ; Reichenow, 1929, p. 893 ; 1932, 
pp. 35-6 ; 1935, p. 369. 

Young trophozoites with simple papillate epimerite. 
Sporonts free, without epimerite. Not forming syzygies. 
Sporocysts not known. 

Remarks. — The genus was based on U. elliptica Mingazzini, 
1891, and among the characters mentioned were " external 
membrane forms a continuous sac round the animal " and 
" protomerite the more dense." Neither of these characters 
is now insisted upon. Porter (1899) described an unnamed 
septate Gregarine from Rhynchoholus , but Crawley (1903), 
thinking that the part of the animal which Porter took to 
be protomerite plus epimerite was only the epimerite, and that 
the Gregarine was a dicystid form, named it as Doliocystis 
rhyncoholi. Kamm (1922) referred it to Ulivina as a second 
species in that genus. Reichenow (1932, 1935) has given 
an amended definition of the genus and considers Sycia 
inopinata Leger, 1892, as idential with Ulivina elliptica 
Mingazzini. The generic character that the epimerite is sur- 
rounded at its base by a ring-like thickening is based on Leger's 
description of S. inopinata. This feature is not possessed 
by U. elliptica, U. rhynchoboli or by the species described 
below. I therefore regard Ulivina as distinct from Sycia. 

311. Ulivina eunicse Bhatia & Setna. (Fig. 186). 

■fUlivina sp., Bhatia & Setna, 1935, p. 312. 
^Ulivina eunicse, Bhatia & Setna, 1938 {in press). 

Trophozoites elliptical, with the protomerite drawn out hke 
a neck and curved backwards. Epimerite slender, wider 
at its base and tapering into a fine needle. The deutomerite 
full of inclusions. Length of protomerite to total length as 
1 : 4-3 ; width of protomerite to width of deutomerite as 1 : 6-3. 
Nucleus oval and situated in the anterior narrower part of 
the deutomerite. Sporocysts not known. 

Dimensions. — Cephalont 340-4 yu. in length ; epimerite 14-8 /^ 
in length ; protomerite 77-7 /a in length and 11-1 /x in maximum 
width ; deutomerite 247-9 /x in length and 70-3 ft in maximum 



Remarks. — The parasite resembles closely U. 
(Crawley), but the cjiioplasm in the protomerite 
dense than in the deutomerite, and the nucleus 
not spherical as in that species. Also there is 
membrane continuous around the animal. 

Habitat. — Intestine of Eunice siciliensis Grube : 
Islands, Port Blair. 

is not more 
is oval and 
no external 


Fig. 186. — Ulivina eunicse Bh. & Set. 
(After Bhatia and Setna.) 

Genus DEUTEROMERA Bhatia & Setna, 1938. 
Deuteromera, Bhatia & Setna, 1938 (in press). 
Sporonts soUtary. Epimerite subcorneal, apex cup-shaped. 
Protomerite and deutomerite showing incomplete secondary 

312. Deuteromera cleava Bhatia & Setna. (Fig. 187). 

fSeptate Gregarine, Bhatia & Setna, 1935, p. 312. 
^Deuteromera cleava, Bhatia & Setna, 1938 (in press). 

Body elongate oval, distinctly septate. Epimerite simple. 



elongated and subconical, with its apex somewhat cup-shaped, 
marked with distinct epicytal striations. Protomerite not 
more densely granular than the deutomerite, broadest at its 
base, width usually greater than its length, showing a somewhat 
obliquely running furrow. Protomerite, together with the 
epimerite, bent at an angle on the deutomerite. Deutomerite 
widest anteriorly, and narrower and rounded posteriorly, 
its maximum width may be somewhat greater than the length ; 
marked by two incomplete septa running inwards from one 
side. Length of the protomerite to total length as 1 : 3*8, 
width of the protomerite to width of the deutomerite as 
1 : 1-3. Nucleus large, oval, situated near the posterior end, 
containing a single large, spherical, eccentrically placed karyo- 
some, surrounded by a narrow clear area and densely packed 

Fig. 187. — Deuterotnera cleava Bh. & Set. 
(After Bhatia and Setna.) 

chromatin granules. Sporont solitary, proportionately 
narrower and much more elongated than the cephalont ; 
may be bent upon itself, and the deutomerite may show 
incomplete septa. Nucleus in the sporont may be oval or 
spherical, but is similar in structure to that in the cephalont. 
Cyst- and spore-formation not known. 

Dimensions. — Cephalont up to 402-5 ju, in total length 
epimerite 105 ju, in length and 17 [x in maximum width 
protomerite 105 /x in length and 147 /^ in maximum Avidth 
deutomerite 192-5 /x in length and 203 /x in maximum width, 
nucleus 52-5 /u by 45-5 /x. Sporont up to 420 /x in total length 
and 105jLi in maximum width, nucleus 42 /x in diameter. 

Remarks. — The furrow on the protomerite and the incom- 
plete septa and a furrow in the deutomerite strongly recall 


the more complete segmentation of the posterior part of the 
deutomerite in Metamera schubergi Duke, 1910, known from 
the intestine of certain leeches, and the segmentation of both 
protomerite and deutomerite in Tseniocystis legeri Cognetti, 
1911, from the coelom of an oligochaete, but there are various 
important differences. 

Habitat. — Intestine of Eunice siciliensis Grube : Andaman 
Islands, Port Blair. 

Genus CONTORTIOCORPA Bhatia & Setna, 1935. 

Contortiocorpa, Bhatia & Setna, 1935, p. 312 ; 1938 (in press). 
Sporont sohtary. Body spirally twisted upon itself. 

313. Contortiocorpa prashadi Bhatia & Setna. (Fig. 188.) 

"fContortiocorpa prashadi, 1935, p. 312 ; 1938 {in press). 

Sporont sohtary. Body elongate oval, broadest at about 
one-third the length of the body from the anterior end, rapidly 

Fig. 188. — Contortiocorpa prasliadi Bh. & Set. 
(After Bhatia and Setna.) 

narrowing behind the middle, and posteriorly drawn out into 
a narrow rounded tip. It is generally twisted upon itself, 
presenting a number of turns of a spiral and a number of 
marginal projections where the spiral is turning round to run 
over the other surface. Nucleus oval or spherical, with 
a single karyosome, generally situated in the anterior half of 
the body. Some individuals are also found in an untwisted 
or only partially twisted condition. 

Dimensions. — Length of the body in a twisted individual 
318-5 ju., maximum width ^l'?>ii, diameter of the nucleus 37-5 ju,. 
A partially twisted individual measured 398-4 /x, mth a maxi- 
mum width of 170-2 /i,. 

Remarks. — In addition to the typical twisted individual 
there are in the preparations a number of partially twisted or 
untwisted individuals which are beheved to be of the same 
species. One such individual shows an indication of a narrow 
protomerite and a retracted bluntly conical epimerite. Typical 


cephalonts have not been met with, nor have we come across 
any cysts or spores which may be definitely assigned to this 

Habitat. — Intestine of Eunice siciliensis Grube : Andaman 
Islands, Port Blair. 

Family SELENIDIID^ BrasH, 1907. 

The family includes a single genus. 

Genus SELENIDIUM Giard, 1884, emend. 
Brasil, 1907, and Ray, 1930. 

Selenidium, Giard, 1884, p. 192. 

Polyrhabdina, Labbe, 1899, p. 48. 

Selenidium, Caullery & Mesnil, 1899, pp. 80-99 ; Minchin, 1903, 
p. 204 ; Brasil, 1907, pp. 370-97 ; Brasil & Fanthani, 1907, 
p. 518; Wenyon, 1926, pp. 1127, 1136-8; Reicheuow, 1929, 
pp. 876-7 ; Ray, 1930, pp. 370-98 ; Reichenow, 1932, pp. 22-5 ; 
1935, p. 366. 

Trophozoites elongate, vermiform, very narrow and cyhndri- 
cal or wider and more or less flattened, with longitudinal 
myonemes along the entire length of the body. The anterior 
end of the body is provided with a small knob-like organ of 
fixation, and usually contains characteristic chromatic 
bodies. Schizogony, where known, takes place during the 
intracellular condition of the parasite. Gametocyst, where 
known, contains many oocysts, each containing either four 
or eight sporozoites. 

Remarks. — Ray (1930) has re-studied several imperfectly 
known species of this genus and described several new ones. 
He has shown that intracellular schizogony does not normally 
occur in the majority of species studied by him, and is, in fact, 
known to occur in two species only. He lays stress on the 
occurrence, in all the species examined and at all stages of 
their development, of characteristic chromatic bodies at the 
anterior end of the animal. These are usually thread-like, 
sometimes club-shaped, but always of a definite type and length 
in any particular species, and usually fairly constant in number. 
Very little is known about sporogony in the species of this 
genus. In two species gametocytes were seen by Caullery 
and Mesnil in association in the gut, and according to them the 
attachment was by their anterior ends. Ray found that in 
the species examined by him the associates become attached 
by their posterior ends. Spores had been previously seen in 
one species only, and were known to contain four sporozoites. 
Ray found gametocysts and spores in two species, and the 
spore contained four sporozoites in one species and eight 



in the other. In view of the above-mentioned considerations, 
he came to the conclusion that the genus requires drastic 
revision, and will probably have to be dismembered. He also 
supports the view, previously held by Mesnil (1899) and 
Keilin (1923), that the Schizogbegarinaria are a heterogeneous 
and artificial group, and that certain genera now placed therein 
will some dav be transferred to the Eugbegarinaria. 

314. Selenidium amphinomi Bhatia & Setna. (Fig. 189.) 

"fNeniatocystis sp., Bhatia & Setna, 1935, p. 312. 

"f Selenidium amphinomi, Bhatia & Setna, 1938 {in press). 

Trophozoites elongate, vermiform, wider anteriorly, and 
narrower and tapering posteriorly. Anterior end provided 

Fig. 189. 

-Selenidium amphinomi Bh. & Set. 
(After Bhatia and Setna.) 

with a conical knob-hke projection which usually does not 
show the chromatic bodies. The body is more or less circular 
in transverse section and the surface is marked by about 
sixteen longitudinal striations. Nucleus spherical or sub 
spherical, situated in the broader anterior part of the body, 
with a large central karyosome . Larger individuals, apparently 
gametocytes, show a somewhat flattened body, wdth the 
anterior end drawn in but still differing in its appearance 
from the rest of the body, and the posterior end is wider than 
in the younger trophozoites and narrows gradually to a 
SPOK. 2 c 


point. The myoneme striations are more numerous and may 
be about twenty in number. The nucleus in these specimens 
is subspherical or oval, and is placed with its long axis along" 
the length of the body. Association of the individuals is by 
their anterior ends. Gametocysts are subspherical or oval. 
Spore-formation was not observed. 

Dimensions. — Trophozoites 126-8-253-6/i, in length, with 
a maximum width ranging from 10 to 25-3ja ; epimerite 4-7- 
9'5/x. in length and about the same in its width. Gametocysts 
are 47-l-81-6;u, in length by 45-5-62-8/x in width. 

Remarks. — ^Bhatia and Setna (1938) have not found any 
evidence of schizogony in this species, but their observations 
are based on the examination of smears only. The chromatic 
threads or bodies, on the occurrence of which during all stages 
Ray (1930) lays so much stress, were not found in many 
specimens. One specimen, however, showed a single deeply- 
stained thread and two other specimens showed a varying 
number of chromatic dot-like bodies. The association of the 
individuals was found to be by their anterior ends, as was 
described by Caullery and Mesnil (1899) in Selenidium echi- 
natum, Caullery & Mesnil, and contrary to what Ray (1930) 
found in other species. A few gametocysts were encountered, 
but none contained ripe oocysts. 

Habitat. — Coelomic cavity of Amphinome rostrata (Pallas) : 
taken off Port Blair, Andaman Islands. 

Family EMERIID^ Leger. 

Genus EIMERIA Aime Schneider, 1875. 
(Pages 173-97.) 

[106. Eimeria gupti, nom. nov. 

"fEimeria cylindrica, Ray & Das-Gupta, 1936 a, p. 345. 

A new species was recorded by Ray and Das-Gupta from 
Natrix piscator and named E. cylindrica (see p. 179). As the 
name is pre-occupied for E. cylindrica Wilson from cattle, 
the species found by Ray and Das-Gupta is re-named 
E. gupti.] 

315. Eimeria arloingi (Marotel). 

Coccidium arloingi, Marotel, 1905, p. 52. 

Eimeria arloingi, Martin, 1907, p. 6 ; Stevenson, 1911, pp. 355—9, 

pis. xix, XX ; Speigl, 1919, p. 451 ; Velu, 1919, pp. 298-301 ; 

Lerche, 1921, pp. 380-99, pi. xviii ; Schein, 1921, pp. 380-2. 


Eimeria faurei, NoUer, Schiirjohann, & Vorbrodt, 1922, 15 pp., 
7 figs. ; Wenyon, 1926, pp. 843-4 ; Knowles, 1928, p. 364 ; 
Reichenow, 1929, pp. 945-6. 

Eimeria arloingi, Reichenow, 1935, p. 374. 
"f Eimeria arloingi, Taylor, 1938, p. 42. 

Oocysts measure 21-33 ju- by 16-5-22-5jU.. 

Remarks. — ^NoUer, Schurjohann, and Vorbrodt (1922), as 
a result of cross-infection experiments with the Coccidia of 
goats and sheep, established the identity of E. arloingi of 
goats with E. faurei of sheep, and since then the two have been 
regarded as identical. Taylor (1938) has recently reported 
that two strains of Eimeria, viz., E. arloingi and E. faurei, 
have been isolated from local goats at Muktesar , and experiments 
are being conducted to test the pathogenesis of these two 
species in goats and sheep. A strain of these parasites is 
claimed to have been estabHshed by feeding sporulated 
oocysts to goats and sheep. 

E. arloingi is considered as highly pathogenic and was 
recently found in an outbreak among goats and sheep at 
Etah breeding farm. 

Habitat. — Faeces of goats and sheep : United Provinces, 
Muktesar, Etah. 

316. Eimeria cylindrica Wilson. 

Eimeria cylindrica, Wilson, 1931, pp. 1—44, 7 pis. 
iEimeria cylindrica, Taylor, 1938, p. 42. 

Oocysts approximately cylindrical, with uniformly thick 
walls, 19-4-26-8/x in length by 11-9-14-9/z, in width, average 
23-3|a by 13-3 fjL. Ratio of length to width 1-65. 

Habitat. — Faeces of hill bulls : United Provinces, 

317. Eimeria lablbeana Pinto. 

Eimeria pfeifferi, Labbe, 1896, p. 539 ; 1899, p. 59 ; Nieschulz, 
1921, pp. 71-82, pi. iv ; 1925, pp. 479-94, pis. xvii-xx ; Wenyon, 
1926, pp. 857-8 ; NoUer, 1928, p. 195. 

Eimeria labbeana, Pinto, 1928, pp. 1564—5. 

Ei'ineria pfeijferi, Reichenow, 1929, p. 948. 
^Eimeria labbeana, Taylor, 1938, p. 42. 

Oocysts spherical, without a micropyle, and without 
a residual body. Sporocysts with one end pointed and the 
other end rounded, with a refractile knob at the pointed end. 
The sporocyst contains a large residuum. 

Dimensions. — Oocysts measure 15-26|U, in length by 14-24/a 
in breadth. Sporocysts measure on an average 12-5^ by 




Remarks. — Nieschulz (1921) at first failed to infect chicks 
with the Coccidium of the pigeon, but later (1925) succeeded 
in doing so. Wenyon (1926) doubts if E. pfeijferi (Labbe) 
of the pigeon is distinct from E. avium (Rivolta & Silvestrini) 
of the chick. Pinto (1928) has proposed a new name, E. lab- 
beana, for Coccidium pfeijferi Labbe and Eimeria pfeifferi 

Genus PROTEOSOMA Labbe, 1894. 

(Pages 247-59.) 

318. Proteosoma passeritse (de Mello & da Fonseca). (Fig. 190.) 

^Plasmodium passeritae, de Mello & da Fonseca, 1938, pp. 47-8, pi. vi. 

Ring-forms with the cytoplasm very compact or more or 
less vacuolated, with small dots of pigment scattered all over 
the bod3^ Schizonts minute, pyriform or amoeboid, devoid 

Pig. 190. — Proteosoma passeritae (de Mello & da Fonseca). A, B, ring 
forms ; G, D, schizonts ; E, cruciform rosette. (After 
de Mello and da Fonseca.) 

of pigment, which may even be absent in larger roundish 
forms. Chromatin divides into two, three or four granules, 
thus determining the number of resulting merozoites, which 
may be arranged in a cross-hke manner. Infected red corpuscles 
do not show any alteration. 

Dimensions. — Ring-forms 1-3 /x, rosettes 3-4 /x. 

Remarks. — ^Apart from the fact that the rings are pigmented, 
the resemblance to Babesia qvudrigemina (NicoUe) is very 
striking, as is also the case with P. minasense (Carini & Rudolph) 
from certain lizards. 

Habitat. — Blood of the green snake, Passerita mycterizans 
Daud. : Portuguese India, Nova Goa. 


Genus ANAPLASMA Theiler, 1910. 
(Pages 325-6.) 
319. Anaplasma marginale Theiler. 

" Marginal points," Smith & Kilborne, 1893. 

Anaplasma marginale, Theiler, 1910, pp. 135—7 ; Wenyon, 1926^ 
pp. 1053-6. 
"f Anaplasma sp.. Ware, 1932, pp. 31—2. 
^Anaplasma m,arginale, Ware, 1938, p. 212. 

Spherical granule, varying in size from 0-1 to 0-5 /x ; staining 
bright red with Romanowsky's stain, and situated near the 
margin of the red blood-corpuscle. 

Remarks. — ^Anaplasmata have been encountered in the 
blood of hill bulls at Muktesar and the animals were regarded 
as " carriers." Dias and Aragao (1914) claimed to have 
brought about the production of these bodies in cattle by the 
simple injection of certain poisonous substances such as trypan 
blue. Du Toit (1928) expressed the view that the production 
of true Anaplasmata in cattle by repeated injection of trypan 
blue can only be brought about when the cattle are " carriers," 
the injections causing a breakdown in their immunity, with 
the appearance of parasites in their blood. Ware (1932) 
reported the appearance of these parasites in hill bulls after 
a series of trypan blue injections. 

Habitat. — Blood of cattle : United Provinces, Muktesar ; 
Central Provinces. 

Genus BERTARELLIA Carini, 1930. 

(Pages 326-7.) 

320 Bertarellia carinii de Meja-elles. 

Bertarellia carinii, de Meyrelles, 1938, pp. 49-53, figs. 1 & 2. 

Usually roundish or elliptical, rarely oval, pyriform or ring- 
like bodies stained violet with Leishman's or May-Grunwald 
Giemsa's stain. Each such chromatic point or anaplasmoid 
body is surrounded by a clear halo, often difficult to dis- 
tinguish, and roundish or elliptical in form. The parasite 
may be central, polar or peripheral in position in the red 
blood-cells, or may even be found free in the plasma. The 
red cell may contain more than one parasite, and the halo- 
like portion may contain two or even three chromatic granules, 
suggesting binary division. 

Remarks. — The organism differs from B. calotis, as the 
cytoplasm is never stained blue as in that species. It is, 
however, identical with the parasite found in blood-films 
from two Brazihan tortoises. 

Habitat. — Blood of the Indian tortoise, Lissemys punctata 
granosa Smith : Portuguese India, Nova Goa. 



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